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From Middle to Upper Class Sprawl? Land Use Controls

and Changing Patterns of Real Estate Development in
Northern New Jersey
a a b a
Thomas K. Rudel , Karen O’Neill , Paul Gottlieb , Melanie McDermott & Colleen
c
Hatfield
a
Department of Human Ecology, Rutgers University
b
Department of Agricultural, Food, and Resource Economics, Rutgers University
c
Biological Sciences, State University of California–Chico
Version of record first published: 28 Mar 2011.

To cite this article: Thomas K. Rudel , Karen O’Neill , Paul Gottlieb , Melanie McDermott & Colleen Hatfield (2011): From
Middle to Upper Class Sprawl? Land Use Controls and Changing Patterns of Real Estate Development in Northern New Jersey,
Annals of the Association of American Geographers, 101:3, 609-624

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 From Middle to Upper Class Sprawl? Land Use
Controls and Changing Patterns of Real Estate
Development in Northern New Jersey
Thomas K. Rudel,∗ Karen O’Neill,∗ Paul Gottlieb,† Melanie McDermott,∗ and Colleen Hatfield‡
∗
Department of Human Ecology, Rutgers University
†
Department of Agricultural, Food, and Resource Economics, Rutgers University
‡
Biological Sciences, State University of California–Chico

During the past twenty-five years the land use controls that shape residential real estate development in the
United States have changed in potentially significant ways. From the 1950s to the 1980s, land use laws promoted
middle-class sprawl by reserving extensive tracts of land for the construction of moderately priced, single-family
Downloaded by [The University of British Columbia] at 21:15 03 February 2013

homes on lots of less than one acre. More recently, suburbs have adopted land use controls that promote upper
class sprawl by reserving large areas for the construction of small numbers of expensive homes on spacious lots.
This regulatory shift can be explained in several ways: a homevoter hypothesis that derives the new controls from
the economic interests of suburban homeowners and a regional spillover hypothesis that attributes the adoption
of new controls to desires by planning commissioners, consultants, and nongovernmental organizations to do as
other communities are doing. We assess these explanations through a case study of changing land use controls in
the suburban New Jersey Highlands west of New York City. Between 1975 and 2002 the region saw large increases
in preserved open space, a doubling of the required minimum lot area for houses, increases in the real price of
housing, declines in the number of newly constructed homes, and a shift in residential real estate development
toward the urban core. Multivariate analyses of the changes in land use controls support the regional spillover
hypothesis. The implications of this dynamic for conservation policies, environmental injustices, and greenhouse
gas emissions are briefly explored. Key Words: down zoning, land use controls, open space, sprawl.

Durante los pasados veinticinco años los controles de uso del suelo que configuran el desarrollo de la propiedad
raı́z residencial de los Estados Unidos se han transformado de maneras potencialmente significativas. De los
años 1950 a los 1980, las leyes de uso del suelo promovieron un desparramamiento de la clase media al reservar
extensiones de tierra muy grandes para la construcción de casas unifamiliares de precio moderado sobre lotes de
menos de un acre. En tiempo más reciente, los suburbios han adoptados controles de uso del suelo que promueven
una dispersión similar de clase alta al reservar grandes áreas para la construcción de un número pequeño de
casas lujosas en lotes espaciosos. Este cambio regulatorio puede explicarse de varias maneras: una hipótesis del
votante de hogar que deriva los nuevos controles de los intereses económicos de propietarios de casas suburbanas,
y una hipótesis de excedentes regionales que atribuye la adopción de nuevos controles a los caprichos de
los comisionados de planificación, consultores y organizaciones no gubernamentales por imitar lo que otras
comunidades están haciendo. Evaluamos estas explicaciones por medio de un estudio de casos de cambio de
los controles de uso del suelo en el suburbio de las Highlands de Nueva Jersey, al oeste de la ciudad de Nueva York.

Annals of the Association of American Geographers, 101(3) 2011, pp. 609–624  C 2011 by Association of American Geographers
Initial submission, January 2009; revised submissions, March and June 2010; final acceptance, July 2010
Published by Taylor & Francis, LLC.
 610 Rudel et al.

Entre 1975 y 2002 la región experimentó grandes incrementos de espacio abierto preservado, el doble del mı́nimo
requerido para el área del lote para casas, aumentos en el precio real para vivienda, declinación en el número
de nuevas casas construidas y un cambio en el desarrollo de la finca raı́z residencial hacia el centro urbano.
Los análisis multivariados de los cambios en los controles de uso del suelo apoyan la hipótesis de los excedentes
regionales. Se exploran brevemente las implicaciones de esta dinámica en las polı́ticas de conservación, injusticias
ambientales y emisiones de gases de invernadero. Palabras clave: re-zonificación, controles de uso del suelo, espacio
abierto, desparramamiento urbano.

into an urban land use. Farmlands, forest lands, and

O
ne word, sprawl, captures the most salient
change in the North American settlement pat-
tern over the past sixty-five years. Sprawl has
been as controversial as it has been ubiquitous. For al-
most four decades land use planners and land change
scientists have debated the social, economic, and envi-
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wetlands can all be preserved open space in New Jer-
sey. The age of the housing stock and the mix of land
uses define the different types of communities in the
New York metropolitan area. Center cities and adjacent
communities that became built up before World War

ronmental costs of sprawl (Real Estate Research Corpo-
ration 1974; Burchell et al. 2005). Many observers have
decried the higher economic costs, additional energy
expenditures, and environmental injustices associated
with low-density suburbanization (Ewing 1997; Bullard,
Johnson, and Torres 2000; Duany, Plater-Zyberk, and
Speck 2001), whereas others have defended sprawl as
the settlement pattern most preferred by Americans
(Audirac, Shermyen, and Smith 1990; Gordon and
Richardson 1997; Bruegmann 2005). Land use controls
figure centrally in this debate because they prescribe
sprawl and, in so doing, contribute to the problems that
come with it.
Most recently, the middle-class sprawl of the 1950s
and 1960s seems to have given way, after changes in
land use laws, to a post millennial, upper class sprawl
in some metropolitan areas. Builders in these places
stopped building small single-family homes on lots of
less than one acre and started building larger single-
family homes in more spacious subdivisions. In this
context it seems important to understand the causes
for the changes in land use controls that have reshaped
suburban sprawl and the effects of these changes in
controls on the building industry. We investigate these
issues through a study of changing patterns of land use
control and real estate development in the New Jersey
Highlands west of New York City.
Definitions of recurring concepts will help to clarify
the following analysis. Exclusionary zoning prohibits all
urban land uses except large-lot residential real estate
development over large areas of a community. If these
ordinances allow other land uses, they are confined to a
very small area of the community. Required minimum
lot area refers to the legal requirement in zoning ordi-
nances that each new housing unit be built on a certain
minimum amount of land. Preserved open space makes
it impossible to convert an undeveloped parcel of land
II constitute the urban core. Low-density residential
land uses predominate in the suburbs constructed after
World War II. A mix of working farms, wealthy resi-
dents on estates, and middle-class commuters character-
ize the exurbs. We use the terms exurbs and rural–urban
fringe communities interchangeably. The term sprawl
has multiple meanings (Fulton et al. 2001; Hasse and
Lathrop 2003; Torrens 2006). Because this article fo-
cuses on land use controls that shape sprawl by con-
trolling the density and location of buildings, we use
a definition of sprawl that emphasizes the density di-
mension. Sprawl refers to low-density residential devel-
opment that often occurs along the rural–urban fringe
of metropolitan areas. Upper class sprawl features es-
tates and large-lot subdivisions of single-family homes.1
In the particular context of postmillennium northern
New Jersey, new homes in these landscapes usually
have at least two acres of land and cost more than
$800,000.2
Theories of Change in Land Use Controls
Efforts to alter local land use controls almost always
begin with a developer’s proposal to build houses on
a particular tract of land. Under these circumstances
the opponents of a proposal can limit real estate
development by preserving open space, usually through
purchase of the land and its transfer to a government
or a land trust that agrees not to develop the land
(Whittaker 1999). Alternatively, residents in a
community can lobby for a change in the land use
regulations that reduces the number of homes that can
be built on a tract of land by increasing the minimum
lot area required for each new housing unit. The latter
technique is often referred to as down zoning (Adelaja
and Gottlieb 2009).
 From Middle to Upper Class Sprawl?
Although different in their ecological effects and
in the ways that they restrict development, preserved
open space and increases in required minimum lot areas
both raise the prices of nearby houses. Agreements to
preserve open space usually result from lengthy negoti-
ations in which neighboring property owners, environ-
mental nongovernmental organizations (NGOs; e.g.,
land trusts), public officials, and developers all partici-
pate after the developer has proposed to build on a tract
of land. To coopt the opposition, developers sometimes
agree to donate a portion of a contested tract of land
to the township as open space in return for approval
to build houses on the remaining land (Beuschel and
Rudel 2010). Limited to building a smaller number of
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homes on spacious lots, developers respond by building
larger homes that generate more profits per unit.
Increases in required minimum lot areas have some-
what different origins. Again, visible signs of increasing
pressure from developers, in the form of proposals to
build increased numbers of homes, provoke a reaction
among local residents and planners. The heads of home-
owner associations and planning commission members,
with legitimating support from planning consultants,
typically push for larger minimum lot areas to limit
the scale of future real estate developments and, in so
doing, preserve visually pleasing environmental ameni-
ties and a vaguely defined “country” way of life (Rudel
1989; Duncan and Duncan 2004). If some communi-
ties in a region have recently increased their required
minimum lot areas, the impetus to do likewise grows
among planners in a region’s other communities. Cur-
rent homeowners usually endorse these changes in land
use controls because the expanded open space and down
zoned districts restrict the supply of new homes, thereby
ensuring that house prices continue to rise and the po-
sitional good of residence in an exclusive community
retains its value.
These accounts of change in land use controls feature
either homeowners or planners as agents of change.
Change in land use controls could conceivably follow
either one or both of these paths. We elaborate on these
homeowner-driven and planner-promoted processes of
change next.
The Homevoter Hypothesis
In one theory about how change in land use controls
occurs, referred to as the homevoter hypothesis (Fischel
2001), homeowners compute the effects of additional
real estate development on the value of their homes and
act accordingly, opposing proposed developments that
611
threaten to reduce the value of their homes and allowing
developments that promise to raise the value of their
homes. Following this line of thinking, homeowners in
high-status communities would reject all but the most
expensive new real estate developments. Homeowners
in nearby, but less exclusive, suburbs would be more
tolerant of new developments because a larger fraction
of them would raise the status of their neighborhoods
and the value of their homes.
The historical dynamics between real estate devel-
opers and suburban homeowners during the twentieth
century provide some support for this line of reason-
ing. Beginning in the mid-1950s, the construction of
interstate highways made many communities on the
outskirts of center cities more accessible by car. Coali-
tions of local businessmen and politicians formed to
promote and build homes on the newly accessible lands
outside of cities (Molotch 1976; Rudel 2009). Their
successes in home building set in motion changes that
eventually make it more difficult to continue building
homes in these places. The new houses passed, through
purchase, into the hands of homeowners who worked
in subsequent decades to prevent the construction of
additional homes in these communities (Rudel 1989;
Furuseth and Lapping 1999; Hogan 2003; Duncan and
Duncan 2004). As in processes of succession (Smith
1974), real estate developers and their allies created
over time the conditions for their own demise, at least
in some places.
The evidence of homeowner activism is abundant.
As soon as entrepreneurs built commuter railroads in
the mid-ninteenth century, upper class residents of
American cities moved to the new suburbs along the
rail lines and began devising ways of preventing poor
people from settling nearby (Jackson 1985). Well-to-do
people pressed for limits on the number and status of
people who could live near them in part because places
of residence had become positional goods with value
that rested in large part on their exclusivity (Hirsch
1976). These places had value for the owner because
most people could not live there. To be sure, places of
residences provided material benefits like shelter, but
they also provided status with their manicured lawns
and well-regarded schools (Robbins 2007). The status
associated with an attractive suburban community
became tangible for residents when they sold their
homes. Homes in high-status communities sold for
substantially more than they would have in lower
status communities. Under these circumstances the
current residents’ desire to maintain the exclusive
status of their community often conflicted with the
 612 Rudel et al.
developers’ desire to build more homes nearby. New
homes, if modestly priced, could dilute the value of
a residence as a positional good because they call
into question the exclusive status of the community.
Environmental concerns mix with questions about
status. The new real estate developments arouse oppo-
sition from environmentalists because they transform
landscapes, threatening local biodiversity and polluting
water.
Status concerns figure centrally in these accounts
of expanding settlements because the new real estate
developments stratify places as they grow, creating
middle-class suburban centers in some places and more
exclusive upper class residential communities in other
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places (Logan 1978; Rudel 1989). Sometimes these
stratifying processes have racial and ethnic implica-
tions. At the onset of suburbanization after World War
II, rural–urban fringe communities contained hetero-
geneous land uses: estates in one district, large farms
in another area, and landfills in a third place. Seeking
inexpensive housing in the suburbs, small communities
of African Americans often located in the places with
landfills that typically had more permissive land use
controls (Bullard 2000; Wiese 2004). Some of these
places went on to become populous, middle-income
“edge cities” (Garreau 1992) or the new centers of
polycentric metropolitan areas (Torrens 2006). An
awareness of these stratifying processes among impor-
tant actors (planners and citizens) usually increases
their commitment to stopping further development.
Previous studies of land use controls also provide
empirical support for this line of reasoning. Exclu-
sionary zoning occurs primarily in upper class suburbs
(Baldassare and Protash 1982; Protash and Baldassare
1983; Rudel 1989; Logan and Zhou 1990; Rothwell
and Massey 2008), and these regulatory tools raise the
price of housing (Glaeser 2002; Quigley and Rosenthal
2005). If changes in land use controls conform to this
dynamic, we would expect the following patterns of
land use control and conversion:
1. The suburbs with the highest socioeconomic status
would enact the most restrictive land use controls.
2. Most real estate development would continue to oc-
cur in suburbs, but it would occur primarily in the
lower status communities with more permissive land
use controls.
3. Developers would also build farther out from the city
center in rural–urban fringe communities that con-
tain larger tracts of less expensive land (von Thunen
[1826] 1966) and more permissive land use controls.
First-home purchasers would then learn to “drive un-
til you qualify,” looking for a home farther from the
city center where the reduced price of housing would
enable them to qualify for a mortgage (Burchell
et al. 2005).
The Regional Spillover Hypothesis
An alternative pattern, dubbed the regional spillover
hypothesis, would lead to a more uniform, exclusion-
ary pattern of land use controls across communities in
suburban regions. Communities would adopt more re-
strictive land use controls (either preserved open space
or increased minimum lot areas) when they see others
in the region making similar changes (DiMaggio and
Powell 1983). A growing network of for-profit firms
(planning consultants) and nonprofit organizations
(regional coalitions of community organizations and
environmental NGOs like the Nature Conservancy)
provide local groups with information about initiatives
in other communities, advice about legal and political
strategies, and financial assistance in preserving tracts
of land (Botshon 2007).
This line of argument generates five expectations
about land use conversion and control in the region.
1. Because differences in land use controls between
communities would be relatively small, variations in
socioeconomic status would not explain the patterns
of change in land use controls.
2. Because new ideas in planning have in recent
years stressed the importance of ecological zoning
(Honachefsky 1999) that restricts building on rugged
tracts of land, topography would predict changes in
land use controls.
3. With relatively uniform increases in land use restric-
tions across the region, the amount of new residential
construction would plummet, with implications for
the larger regional pattern of residential building.
4. The drive until you qualify effect would not be very
apparent in building statistics because the spread of
requirements for large minimum lot areas would ex-
hibit isomorphic tendencies, extending beyond the
wealthiest communities to the exurban, rural–urban
fringe communities far from the urban center.
5. Finally, more residential real estate development
would occur in the urban core.
This last expectation requires a more detailed
explanation. Small-scale developers can respond to
more restrictive land use controls by building smaller
 From Middle to Upper Class Sprawl? 613

numbers of more expensive homes, but large-scale

developers find it more difficult to build under these
circumstances without changing their business model
in fundamental ways. Given the large scale of their
building projects, big developers must regularly locate,
acquire, and then develop large tracts of land (Torrens
2006). In a context where land use controls place many
restrictions on real estate developers, land assembly
becomes a tremendous challenge for them (Rudel
2009). The earlier development of single-family homes
on some tracts of land, coupled with the designation of
other tracts of land as open space, significantly reduces
the supply of undeveloped land. If the remaining un-
developed land exists in fragments, the land assembly
Downloaded by [The University of British Columbia] at 21:15 03 February 2013

problem becomes even more severe, and builders might

not be able to acquire enough of it in any one place
to construct a large-scale development. The physical
features of the land become important under these
circumstances. If a region contains significant amounts
of steeply sloped, unbuildable land, the difficulties of
land assembly multiply even further. Some low-density
suburbs become “built out”; in other words, real estate
developers can no longer find tracts of land in these
communities that they can develop (Conway and Lath-
rop 2005). Under these circumstances developers might
look to build in the older urban core where the higher
permitted densities in the land use regulations make
it possible to build developments with hundreds of
smaller, less expensive units on relatively small tracts of
land.
Although the homevoter and regional spillover
hypotheses generate different expectations about the
dynamics of change in land use controls, the two
hypotheses are not mutually exclusive. They occur
at different spatial scales and feature different agents
of change. The homevoter hypothesis focuses on Figure 1. The New Jersey Highlands.
political processes within communities and identi-
fies the affluent homeowner as the agent of change. The Context
The regional spillover hypothesis describes dynam-
ics that unfold across communities and identifies The geographic focus for this study, the Highlands,
organizations like planning commissions, consulting is a rural–urban fringe and suburban area situated less
firms, and environmental NGOs as the agents of than a two-hour drive west of New York City (see
change. Figure 1). It encompasses approximately 1.5 million
We assess these differing expectations about the ge- acres of Appalachian ridges and valleys, stretching from
ographical dimensions of changes in land use controls the Lower Hudson River Valley along the New York–
through a case study of these changes and real estate New Jersey border to the Delaware River boundary be-
development in northern New Jersey. In the follow- tween New Jersey and Pennsylvania. A series of forested
ing pages we describe the context for the case study, hills and ridges cut by narrow valleys constitute the
the data and methods, the results, and the conclusions backbone of the region. Forests cover more than half of
about an emerging dynamic involving land use controls the land in the region. Oak-hickory forests predominate
and real estate development. in most places, but northern hardwoods, hemlocks, and
 614 Rudel et al.
swamp hardwoods also appear in the region’s forests.
The topography of the southern Highlands is gentler,
and agriculture predominates in this area.
The Highlands provide quality drinking water for
more than 11 million people in the surrounding ar-
eas via ground water aquifers, rivers, and reservoirs
(U.S. Department of Agriculture [USDA] 2002). The
reservoirs provide crucial supplies of water to the re-
gion’s population, particularly during droughts. The
close proximity to the New York City metropolitan
area makes the Highlands a valuable and highly utilized
recreational resource. Approximately 14 million people
visit the Highlands each year to hunt, fish, boat, hike, or
just enjoy the scenery (USDA 2003). The Highlands
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also contains a large portion of the region’s biodiver-
sity. The variety of natural community types, including
freshwater wetlands, swamps and bogs, glades, ravines
and ridges, large contiguous forest tracts, and grasslands
provides important habitat resources for a wide variety
of plants and animals, including 200 plant species and
fifty vertebrate animal species that are listed in invento-
ries for endangered, threatened, or “of concern” species
(USDA 2003).
Although it does not cover a large proportion of
the land, agriculture has been important in sustaining
the region’s economy as well as its rural character. To-
day, approximately 10 percent, or 143,000 acres, of the
Highlands is in crop production. Over 60 percent of the
actively farmed acres are considered to be prime farm
soils, but more than 100,000 acres of lands with prime
soils have already been converted into subdivisions of
single-family homes or commercial buildings (USDA
2003). Both New Jersey and New York have programs
in place to preserve large contiguous blocks of farm-
land to ensure the productivity and economic health of
agriculture over the long term.
Pressure to convert Highland lands to urban uses
increased during the last three decades of the twenti-
eth century. By the late 1960s an unbroken expanse
of built-up land stretched from New York City to the
border of the Highlands, and it began to expand into
the Highlands during the 1970s. Developers converted
about 5,000 Highland acres each year to urban uses
during the 1990s (USDA 2002). The population of the
Highlands increased by 11.5 percent during the 1990s
(USDA 2003). Although this increase is less than the
national average of 13 percent, it is considerably higher
than that observed in either New York (5.5 percent) or
New Jersey (8.9 percent) during the same time period.
We chose the Highlands for study because antisprawl
groups have been active in the region (Solecki, Mason,
and Martin 2004). Their activities exemplify a growing
practice of coordinating land conservation efforts across
organizations and across geographic scales that is evi-
dent in other American places like the highlands east
of San Diego, California (Calthorpe and Fulton 2001;
Hogan 2003; Babbitt 2007; U.S. Bureau of Land Man-
agement 2007) and the Balcones Canyonlands, west of
Austin, Texas (Travis County 2008). For this reason
the findings about changing land use controls in the
Highlands and their effects on regional patterns of real
estate development could have implications for other
metropolitan areas in North America.
Data and Methods
To carry out the following analyses, we developed
measures of change in municipal land use controls. We
focused on municipal land use controls because in New
Jersey, as elsewhere in the northeastern United States,
municipalities have the primary power to regulate land
use. The municipal zoning laws and associated maps
are available for inspection by any citizen, and we used
old zoning maps from earlier eras to chart how land use
controls changed from the mid-1970s to 2002 in each
Highlands community. Because open space preserva-
tion changes the tax status of lands, towns also keep
records about the creation of designated open space
in the community. From these records we constructed
maps that depict the changing extent of preserved open
space in Highlands communities.
Zoning maps for the Highlands municipalities from
2002 were available in a vector-based geographic infor-
mation system (GIS), and they provided a template on
which we mapped the earlier zoning changes and open
space designations in separate layers through on-screen
digitizing in ArcGIS (ESRI 2008). After we created
maps of changes in minimum lot areas and changes in
the extent of preserved open space, we used the GIS
to compute measures of change for each municipal-
ity. We computed the average required minimum lot
area at Time 1 in Town A by calculating the aver-
age required minimum lot area across a community’s
residential zones, weighted by the size of the different
zones (Rudel 1989). A similar procedure gave us the
average minimum lot area at Time 2. From these num-
bers we calculated the change over time in the required
minimum lot area in a community. For preserved open
space, we used open space maps from different points
in time to calculate the extent of increases in open
 From Middle to Upper Class Sprawl? 615

Table 1. Measures for variables in the bivariate and multivariate analyses

Preserved Open Space, Change in % of Municipal Land Area: The extent, computed in a municipal GIS, of preserved open space at Time 2
minus the extent of preserved open space at Time 1. See text for further details.
Minimum Lot Area, Change in Average Required Area: Change in the average minimum lot area, computed in a municipal GIS, at Time 2
minus the average minimum lot area at Time 1. This variable has been logged. See text for further details.
Borough: Borough versus township designation. Source: State of New Jersey (http://www.nj.gov/cgi-bin/infobank/munisearch.pl). Boroughs
are the old commercial centers of many townships. To exert more control over municipal services, they incorporated as separate local
governments.
House Prices: Mean Prices of Sold Homes in the Highlands. Source: Garden State Multiple Listing Archives. House Values, 1970; U.S.
Bureau of the Census, 1971, 1970 Census of Housing. General Housing Characteristics: New Jersey.
Minimum Lot Area, Required for Housing Units, Average: As described in the text, calculated from a GIS made out of municipal zoning
regulations for a particular year.
Median Family Income, 1969: Source: U.S. Bureau of the Census, 1973, 1970 Census of Population. Characteristics of the Population: New
Jersey.
Open Space, Extent: As described in the text, calculated from a GIS constructed from municipal maps and records.
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Population Density, 1970: Source: U.S. Bureau of the Census, 1970 Census of Population. Characteristics of the Population: New Jersey.
Residential Land Area, Growth in, 1995–2002: Land cover data from 1995 Landsat and 2002 Landsat and Spot Satellite Imagery, arrayed in
a USDA Forest Service GIS at the Center for Remote Sensing and Spatial Analysis at Rutgers University. Calculated for municipalities.
Slopes with an Incline > 15 Percent, Percent of Land Area: Calculated from USGS Digital Elevation Model, 30 meter Resolution
(Landsat), 2002. Available in Center for Remote Sensing and Spatial Analysis database.

Note: GIS = geographic information system; USDA = U.S. Department of Agriculture; USGS = U.S. Geological Survey.

space conservation in communities between Time 1 and
Time 2.
To analyze the changing pattern of land use con-
trols, we merged the vector data on land use controls
with data from two other sources. A USDA Forest Ser-
vice study of the Highlands (USDA 2003) assembled
an extensive, raster-based data set on natural resources
in the Highlands and arrayed these data in a GIS. The
data come from a variety of remotely sensed and terres-
trial sources. We converted raster data on steep slopes
(greater than 15 percent) and land cover change to vec-
tor data by computing municipal totals for these land
attributes and adding them to the other vector data.
We also added nonspatial, demographic, and socioeco-
nomic data to the ecological and land use control data
for the municipalities. The measures for these variables
and the two land use control measures, along with their
sources, are briefly described in Table 1.
To identify the forces driving changes in land use
controls, we analyzed variations in the extent of in-
creases in preserved open space and minimum lot areas
across the eighty-three Highlands communities. Table 2
provides a summary of the trends in some of these
variables over time in the Highlands communities. In
the multivariate analyses reported in Table 3, we logged
the required minimum lot area variable to reduce the
influence of outliers. Problems of multicollinearity and
simultaneity bias do not compromise the analyses. The

Table 2. Society and land use controls in the New Jersey Highlands: Trends from the mid-1970s to 2007 for the eighty-three
Highlands communities

Date of measures: Approximate

Variables 1970–1975 1992–1995 2000–2002 2005–2007

Average required minimum lot area (1970s, 2002) 1.49 acres 2.88 acres
% of Highlands land area preserved as open space (1970s, 2002) 7.6% 29.1%
Housing units constructed (1994, 2001, 2007) 3,260 2,611 1,109
Average selling price of homes in unadjusted $ (1992, 2005) $219,889 $412,146
Median family income as a % of U.S. average (1969, 2000) 137.5% 207.9%

Note: The actual dates when the data were collected are listed in parentheses under the variable name. Where it lists a range of dates, like 1970s, it means
that the dates of the data varied from municipality to municipality.
Sources: Average required municipal lot areas: Municipal archives; Open space preserves: Municipal archives; Housing units: U.S. Census Bureau, building
permit data; Average selling prices of homes: Multiple Listing Services, New Jersey; Median family incomes, 1969: United States Census, 1970.
 616 Rudel et al.

Table 3. Preserved open space and large lot zoning, 1975–2002: Spatial regression analyses

(5) Large lot (6) Large lot

(2) Open space (3) Open space zoning with zoning with
(1) Open space, with median family with median house (4) Large median family median house
Variables % of area income, 1969 value, 1970 lot zoning income, 1969 value

Spatial lag 0.270∗ (0.111) 0.281∗ (0.111) 0.271∗ (0.111) 0.137 (0.103) 0.174∗ (0.105) 0.161 (0.105)
Steep slope, > 15% 1.214∗∗∗ (0.267) 1.281∗∗∗ (0.271) 1.255∗∗∗ (0.270) 5.227∗ (2.120) 6.029∗∗ (2.176) 5.812∗∗ (2.162)
Borough (yes/no) −0.109∗∗∗ (0.022) −0.108∗∗∗ (0.021) −0.108∗∗∗ (0.022) −0.340∗ (0.169) −0.354∗ (0.168) −0.347∗ (0.168)
Population density, −0.036∗∗∗ (0.007) −0.035∗∗∗ (0.008) −0.035∗∗∗ (0.007)
1970
Open space, 1970s 0.900∗∗∗ (0.123) 0.870∗∗∗ (0.125) 0.871∗∗∗ (0.128)
Minimum lot area, 0.374∗∗∗ (0.063) 0.356∗∗∗ (0.063) 0.358∗∗∗ (0.064)
1970s
Median family 0.051 (0.044) 0.045 (0.035)
Downloaded by [The University of British Columbia] at 21:15 03 February 2013

income, 1969
Median house 0.011 (0.013) 0.013 (0.010)
value, 1970
R2 0.632 0.635 0.633 0.691 0.698 0.696
N 79 79 79 78 78 78

Note: The top figures in each cell of the table are unstandardized regression coefficients; the numbers beside them, in parentheses, are the associated standard
errors. The units of analysis are municipalities.
∗ p < 0.10.
∗∗ p < 0.01.
∗∗∗ p < 0.001.

variance inflation factors are all below 1.6 and the
condition indexes are all below 15; these levels are well
below the values that indicate the presence of serious
multicollinearity in the data (Kennedy 1985). The
independent variables either do not vary (e.g., land area
in steep slopes) or they are measured at the beginning
of the period under examination, so simultaneity biases
(reciprocal effects between independent and dependent
variables) are not present in the equations. The residu-
als from the equations are normally distributed. Delet-
ing the most influential cases from the equations did
not alter the substantive results presented in Table 3.
Spatial autocorrelation could compromise the value
of the multivariate analyses, with commitments to con-
serve open space and increase minimum lot areas in
one community affecting decisions about open space
and minimum lot areas in neighboring communities
(Rudel 1989; Brueckner 1998). There are modest levels
of global spatial autocorrelation in the data. Moran’s I
was .1737 ( p = 0.02) for changes in open space and
.2124 ( p = 0.01) for changes in minimum lot areas.
To account for these spatial effects, we estimated spa-
tially lagged models. Given our theoretical interest in
the regional spillover hypothesis with its emphasis on
the effects of changes in land use controls in regional
clusters of communities, the spatial lag model of auto-
correlation seems most appropriate because it allows for
feedback effects in which events in neighboring locales
influence later changes in regulations in the commu-
nities of interest. To uncover subregional patterns in
the spillover effects, we looked at local indicators of
spatial association (LISA; Anselin 1995). These results
are presented in the next section.
Results
Municipal land use controls in the New Jersey High-
lands underwent dramatic changes between 1975 and
2002 (Table 2). The amount of preserved open space
increased from 7.6 percent to 29.1 percent of the land
area in the eighty-three Highlands communities be-
tween the 1970s and 2002.3 The increases typically oc-
curred in small increments as local activists, organized
into conservation coalitions, secured funds from public
and private sources to preserve farmlands and forested
tracts of land. The small increments in preserved open
space depicted for Washington Township in Figure 2
illustrate how open space expanded during the three
decades.4 The average minimum lot area required to
build a single-family home in the region expanded from
1.49 acres in 1975 to 2.88 acres in 2002 (Table 2).5
Not surprisingly, housing prices in the region increased
sharply during the latter part of the period. The average
 From Middle to Upper Class Sprawl?
Figure 2. Open space acquisition in
Washington Township, Morris County,
Downloaded by [The University of British Columbia] at 21:15 03 February 2013
New Jersey, 1970–2005. Source: Wash-
ington Township Planning Commission
Maps.
price of a home in the eighty-three communities in-
creased from $219,889 to $412,146 between 1992 and
2005 (Table 2). This increase represents, after adjust-
ing for inflation, a 47.6 percent increase in the real
price for housing in the region over the thirteen-year
period.6 Over time these trends made Highlands res-
idents wealthier than other Americans. In 1970 the
median family income of the Highlands population
was 137.5 percent of the median family income in the
United States. By 2000 the median family income of
Highlands residents had risen to 207.9 percent of the
median for the U.S. population (U.S. Bureau of Census
2010).
The large magnitude of these changes in land use
controls, housing prices, and family income raises ques-
tions about both the agents of change and the effects
of these changes at a larger scale in the building pat-
terns for the entire metropolitan area. Correlational
analyses answer some of these questions. To an appre-
ciable extent, places in the Highlands that conserved
a lot of open space also tended to be municipalities
617
that increased their minimum lot areas (r = 0.399, p <
0.001). Open space preservation and increases in mini-
mum lot areas occurred together because they stemmed
from the same driving force, real estate development.
Increases between 1995 and 2002 in the proportion of
a community’s land developed for single-family homes
associated positively with increases in preserved open
space (r = 0.300, p < 0.01) and minimum lot areas
for new houses (r = 0.217, p < 0.05) across the eighty-
three municipalities. In other words, development pres-
sure and changes in land use controls tended to occur
in the same places.
The spatial regression analyses presented in Table 3
provide a more complete picture of the human ecolog-
ical conjuncture that led to the changes in land use
controls.7 The changes in the extent of open space and
the size of minimum lot areas followed a similar pat-
tern. Boroughs, small, densely settled, and long built-up
municipalities, experienced little change in open space
or minimum lot areas. Municipalities with substantial
amounts of steeply sloped land and abundant amounts
 618
of preserved open space in the 1970s saw considerable
increases in open space during the three-decade pe-
riod. Similarly, municipalities with large amounts of
steeply sloped land and high minimum lot areas in the
1970s experienced the largest increases in minimum
lot areas. Lower population densities in 1970, signify-
ing a relative abundance of undeveloped land, predicted
larger increases in minimum lot areas between the 1970s
and 2002. Interestingly, socioeconomic variables with a
stratification dimension, like median housing prices and
median family incomes in 1970 (in columns 2, 3, 5, and
6 of Table 3) did not distinguish well those Highlands
communities that adopted the most restrictive land use
controls. This pattern, coupled with the significance of
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the steep slopes variable, indicates support for the re-
gional spillover hypothesis rather than the homevoter
hypothesis.
LISA analyses specified the geographic patterns of
change. The most dramatic cluster of increases in open
space occurred in the Highlands communities with
rough topography along the border with New York
State. A cluster of large increases in required mini-
mum lot areas occurred in the transportation corridor
along I-78 (see Figure 3). Large required minimum lot
areas in one community prompted calls in neighbor-
ing communities for increases in their minimum lot
areas (Tewksbury Planning Commission 2003). In this
fashion the increases in minimum lot areas in the I-78
transportation corridor spread westward over time with
development pressure from New York City.
Figure 3. 1970–2002 minimum lot areas in the I-78 corridor: Pat-
terns of association. Dark indicates strong association ( p < 0.01);
gray indicates modest association ( p < 0.05).
Rudel et al.
The magnitude of these changes in land use controls
suggests that they might have had an impact on
real estate development in the larger metropolitan
region. With these changes in mind, we examined
building statistics for residential real estate in northern
New Jersey for the period between 1994 and 2007.
Figure 4 describes a dramatic shift in building patterns
in the larger region. Although the total number
of units constructed increased from 1994–1995 to
2006–2007 by almost 50 percent, from 22,023 to
33,427 units, the location of the new units changed. In
the mid-1990s real estate development concentrated,
as it had for four decades, in suburban communities.
In contrast, in 2006–2007 developers built more
than 50 percent of their units in the urban core
region immediately west of New York City.8 The
numbers of new units constructed declined by 15.7
percent in the suburbs and by 8.5 percent in the
exurbs between 1994 and 2007. The Highlands region,
which straddles the suburban and exurban counties in
Figure 4, appears to have been at the epicenter of the
decline in residential building in the outer reaches of
the metropolitan area. The eighty-three communities
in the Highlands region experienced a dramatic 66.0
percent decline in new housing units constructed, from
3,260 units in 1994 to 1,109 units in 2007. The increase
in residential building in the urban core involved
relatively large developments of condominiums and
apartments, often built near major infrastructure
projects like the newly constructed Hudson–Bergen
light rail line that runs along the west bank of the
Hudson River. Several of the developers in these
urbanized places, like K. Hovnanian Homes, had
previously built only in the suburbs.
Interestingly, these data show little evidence of a
drive until you qualify the effect in which developers,
prevented from building in the now exclusive suburbs,
decide to build on less expensive land farther from the
metropolitan center and first-time home buyers decide
to endure the longer commutes to employment centers
because they can qualify for the smaller mortgages
associated with purchasing less expensive housing. New
residential units in these rural–urban fringe locations,
the two northern New Jersey counties farthest from
New York City, declined from 8.6 percent in 1994–1995
to 5.2 percent in 2006–2007 as a percentage of all new
residential building units in northern New Jersey. The
addition of the three adjacent Pennsylvania counties
(Monroe, Northampton, and Pike) to this rural–urban
fringe area does not alter the overall, regional pattern of
 From Middle to Upper Class Sprawl?
Figure 4. Changing patterns of new res-
Downloaded by [The University of British Columbia] at 21:15 03 February 2013
idential building units in northern New
Jersey, 1994–2007 (% of all new res-
idential units). Source: http://censtats.
census.gov/bldg/bldgprmt.shtml.
change in building. The building in the Pennsylvania
counties, as a proportion of all building in the surveyed
counties, actually fell, as it did in New Jersey’s two
exurban counties, between 1994–1995 and 2006–2007.
At least in this region, the drive until you qualify effect
is not visible. Considered together, these patterns of
new unit construction are broadly consistent with the
regional spillover hypothesis and inconsistent with the
homevoter hypothesis.
Discussion
The findings about changes in land use controls and
building patterns in the eighty-three Highlands com-
munities support a regional spillover interpretation of
events. Rather than a process where each homeowner
calculates his or her economic interest and votes ac-
cordingly, land use controls and associated building
619
seem to change in an almost wave-like pattern in which
contiguous sets of communities modified their land use
controls in similar ways over several decades. Develop-
ers seem to behave in a similar way. The significance
of sloped land in the open space regressions (Table 3)
most likely reflects a propensity by developers to emu-
late one another in donating unbuildable sloped land as
preserved open space in efforts to win approval to build
on other tracts of land (Beuschel and Rudel 2010). It
might also reflect a region-wide decline in undeveloped
but developable land that occurs as the region becomes
more built up. This pattern also reflects the priorities
of people in the conservation coalitions who lead the
fight to preserve lands. Steeply sloped lands in the east-
ern United States typically contain forests, high-quality
water sources, and high levels of biodiversity, in addi-
tion to providing recreational opportunities, so activists
make them a high priority for conservation. By reducing
the supply of land available for development, open space
 620 Rudel et al.
conservation would increase the price of new develop-
ments elsewhere in a community. Price-inflating edge
effects would also exist. The location of houses along
the borders of nature preserves increases the prices of
these homes (Daniels 1999; Geoghegan 2002).
The increases in minimum lot area requirements
follow a pattern similar to the preserved open space
increases except that these changes concentrated in
communities that were less built-up in 1970. Here,
too, the stratification variables (median family income,
house value) did not discriminate effectively between
communities that did or did not adopt more restrictive
controls. The doubling of the minimum lot area require-
ments between the 1970s and 2002 reduced the number
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of units that developers could build and made the
communities more exclusive. Over time the new regula-
tions would create districts characterized by upper class
sprawl. The LISA analyses suggest that spillover effects
from regulatory reforms derive from both ecological and
social circumstances. A cluster of designated open space
in the northwestern corner of the Highlands reflects
the concentration of accentuated terrain in that area.
A cluster of dramatic increases in minimum lot areas in
the I-78 corridor reflects the development pressure radi-
ating outward along the interstate from the urban core.
So who and what has driven these regional pro-
cesses? Planning and zoning consultants who practice
in communities throughout the region have frequently
recommended down zoning (Tewksbury Planning
Commission 2003). Planning commissioners in indi-
vidual communities have also become more aware of
changes in land use regulations elsewhere in the region
through their membership in the Highlands Coalition
(www.highlandscoalition.org), a regional umbrella
organization of sixty nonprofit organizations with inter-
ests in the Highlands. NGOs have played a direct role in
efforts to preserve open space. Local NGOs active in the
Highlands Coalition as well as national environmental
NGOs like Audubon, the Nature Conservancy, and
the Trust for Public Lands have played crucial roles in
these conservation efforts. Public sector organizations
have also contributed. Between 1998 and 2002 many of
the municipalities in the region established local funds
for open space acquisition after the state indicated
that they would match local contributions. In 2003
the state’s voters approved an additional $50 million
for the acquisition of open space in the region. Most
recently, during the spring of 2004, after the period
considered in this study, the state of New Jersey passed
legislation to redirect development away from the most
ecologically sensitive portions of the Highlands.
Recent trends in building permits in northern New
Jersey suggest that these trends in land use controls can
in their aggregate affect regional patterns of real estate
development. To be sure, some historically and region-
ally specific trends like the economic boom on Wall
Street, the changing residential preferences of an aging
population, and a reduced program of highway construc-
tion played some role in producing the dramatic changes
in real estate development observed in Figure 4. Nev-
ertheless, it seems plausible that the changes in land
use controls documented for the Highlands communi-
ties have occurred more broadly in the belt of subur-
ban communities surrounding New York City, and that
these trends have played an important role in displac-
ing real estate development to other places (Carruthers
2003), in this instance to the urban core communities
of northern New Jersey.
The findings in Tables 2 and 3 suggest a new dynamic
in the stratification of places. Affluent homeowners
continue to react negatively to plans for nearby real
estate development and push for changes in land use
controls in ways consistent with the homevoter hypoth-
esis. In so doing, they maintain the exclusivity of their
community, and they contribute to the stratification
of places within their metropolitan area. The success
of the regional spillover hypothesis in explaining the
adoption of restrictive land use controls throughout the
Highlands points to a second stratifying dynamic that is
inconsistent with the homevoter hypothesis. With the
increased interaction between planners and open space
advocates in different communities, the less wealthy
suburban communities begin to adopt land use controls
much like the wealthier suburban communities. Under
these circumstances, stratifying processes appear at a
larger geographical scale within metropolitan areas.
The spread of restrictive land use controls across the
Highlands between 1970 and 2002 contributes to the
emergence of an upper class green belt around a middle-
and lower class urban core in a spatially stratified re-
gional city (Calthorpe and Fulton 2001). In this sense,
when the state of New Jersey created a state-supervised,
regional planning district to limit further real estate
development in the Highlands in 2004, it ratified a
trend in land use controls that was already underway
at the municipal level (New Jersey Department of
Environmental Protection 2010). The two stratifying
processes discussed here could emerge in a historical
sequence. During an early historical period, homevoter
political processes create the institutional infrastruc-
ture that at a later date supports the rapid diffusion
of new norms about land use planning across a region
 From Middle to Upper Class Sprawl?
in a manner consistent with the regional spillover
hypothesis.
Does the dynamic described here apply to other
metropolitan areas in North America? Certainly,
the antidevelopment sentiments of homeowners are
widespread and the growth in the numbers of NGOs
that facilitate the spread of antisprawl measures has oc-
curred in many metropolitan areas (Molotch, Freuden-
burg, and Paulsen 2000; Warner and Molotch 2000;
Duany, Plater-Zyberk, and Speck 2001). Prices of new
homes have risen at accelerated rates since 1960 in all
four census regions of the United States (Dwyer 2007).
Nevertheless, the wave of down-zoning and open
space conservation experienced in the Highlands dur-
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ing the late twentieth century might have depended in
part on an interaction between topography and the de-
gree of political fragmentation in metropolitan areas. In
humid southern and midwestern metropolitan areas like
Nashville, Tennessee or Chicago, Illinois, exclusionary
controls in a context with abundant supplies of build-
able land and fragmented local jurisdictions probably
exacerbate sprawl by forcing developers to jump over
the exclusionary communities and build in areas far-
ther from the center city (Carruthers 2003). When the
exclusionary efforts occur in metropolitan areas like Los
Angeles where arid conditions and mountainous terrain
have limited the supply of buildable land, antisprawl ef-
forts could well limit the supply of buildable land to such
an extent that displaced developers turn back toward
the metropolitan center when they search for places
to build on a large scale. This dynamic would explain
the recent increases in the density of settlement in the
Los Angeles metropolitan area (Lang 2003; Bruegmann
2005). Here, too, green-belt-creating regional plans like
Portland’s growth boundary (Song and Knaap 2004)
and Seattle’s regional land use plan (Calthorpe and
Fulton 2001) might have reinforced a regional trend
toward denser settlement that had already begun.
Conclusions and Implications
It is certainly not surprising that more exclusive forms
of land use control have emerged in the United States
during a period of growing income inequality, but their
repercussions for problems of affordable housing, en-
vironmental injustices, and greenhouse gas emissions
remain poorly understood (Ewing et al. 2007; Andrews
2008). The more restrictive controls have both posi-
tive and negative implications for society and the en-
vironment. First, these controls would probably, over
621
time, reduce per capita greenhouse gas emissions be-
cause the homes of most middle-class people would be
less spacious and their commutes to work would in-
volve fewer and less lengthy car trips. In at least some
limited respects, sprawl for the upper class would pro-
mote smart growth for the middle class (Duany, Plater-
Zyberk, and Speck 2001). Although the more spacious
sprawl would increase emissions per capita among upper
class households, the corresponding declines in emis-
sions among the much more numerous middle-class
households should lead to net declines in emissions.
Second, even as an influx of middle-class people into the
older core communities might rejuvenate community
institutions like public schools and reduce the spatial
concentration of poverty (Burchell et al. 2005), the in-
flux of new residents would also probably raise the price
of housing in the older core communities and put afford-
able housing out of the reach of some poor households.9
In effect these controls would promote gentrification
in the urban core with all of its implications for low-
income residents (Lees, Slater, and Wyly 2008). Finally,
the pattern of upper class sprawl described here would
make more salient an often neglected type of environ-
mental injustice by allowing only the wealthy to enjoy
on a daily basis the environmental services that come
with living in the aesthetically preferred, pastoral land-
scapes that often characterize the outer suburbs (Dun-
can and Duncan 2001; Agyeman, Bullard, and Evans
2003). If the post-2007 economic turmoil strengthens
real estate markets for smaller, less expensive housing
units in the urban core of American metropolitan areas,
these speculations about the changing patterns of real
estate development might become tangible realities.
Acknowledgments
This research was conducted under National Sci-
ence Foundation Grant No. SES 0523309. We deeply
appreciate the agency’s support for this research. We
would also like to thank the many citizens and munici-
pal employees who aided us during the collection of the
field data. The authors benefited greatly from comments
made during presentations of this article at Columbia
University, the University of North Carolina, Rutgers
University, and SUNY—Environmental Science and
Forestry. The anonymous reviewers for the Annals and
the editor made a series of comments that significantly
strengthened the article. Finally, we would like to thank
Vanessa Beuschel Boulier, Caroline Phillipuk, Karen
 622 Rudel et al.
Stein, and Peter Vancura for indispensable research as-
sistance on this project.
Notes
1. This definition of upper class sprawl focuses on residen-
tial rather than commercial land uses because there does
not appear to be a distinct pattern of upper class com-
mercial sprawl.
2. The size of building lots that would distinguish between
upper and middle-class sprawl will vary across housing
markets with the price of land.
3. The dates of the sources for the 1970s land use data, usu-
ally municipal zoning maps, varied from municipality to
Downloaded by [The University of British Columbia] at 21:15 03 February 2013
municipality. For that reason we refer to the initial date
of observation for the open space and required minimum
lot area variables as the 1970s.
4. The categorization of open space preservation by decade
in Figure 2 raises questions about the history of the
preservation efforts. These efforts begin with waves
of real estate development, but subsequent, sometimes
lengthy legal proceedings over proposed land uses make
it difficult to predict when the land will become pre-
served open space.
5. To calculate a region-wide change in minimum lot
area, we computed a region-wide average for each year,
weighting the required lot areas by the size of the com-
munity, so the required lot areas for the larger townships
counted more than the required lot areas for the smaller
boroughs.
6. We calculated these rates using a gross domestic product
price deflator.
7. The models in Table 3 were also run using ordinary least
squares (OLS) techniques. Substantively, there were no
significant differences between the OLS and spatial lag
results.
8. To reduce the influence of sharp annual fluctuations in
housing market conditions, we aggregated the building
unit data over two-year periods: 1994–1995, 2001–2002,
and 2006–2007. The core includes three counties—
Essex, Hudson, and Union—plus the southern munic-
ipalities in Bergen County. The remainder of Bergen
County, Passaic County, Morris County, Somerset
County, Hunterdon County, and Middlesex County
constitute the suburban region. Warren and Sussex
counties represent the rural–urban fringe region. With
the exception of Bergen County, which we split into
an urban core and an outlying suburban district, we
have used county lines to delineate the different housing
markets. Although this procedure has a coarse-grained
quality, it accords with the history of real estate devel-
opment in northern New Jersey, with the core being
developed before World War II, the suburban belt be-
tween 1950 and 1990, and the rural–urban fringe after
1990. It also facilitates the comparison of the large-scale
housing markets depicted in Figure 4. Data sources were
the U.S. Census (http://censtats.census.gov/cgi-bin/)
and http://www.wnjpin.net/OneStopCareerCenter/
LaborMarketInformation/lmi18/hist/bpann94.htm, ac-
cessed in 2008 and 2009.
9. Although it is important to note that upper class sprawl
does nothing to provide affordable housing, it should also
be noted that the earlier period of middle-class sprawl
provided affordable housing only through processes of
residential turnover in the urban core communities in
which middle-class homeowners sold sometimes dilapi-
dated structures to landlords who rented these units to
the poor (Burchell et al. 2005).
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Correspondence: Department of Human Ecology, Rutgers University, 55 Dudley Road, New Brunswick, NJ 08901, e-mail:
rudel@aesop.rutgers.edu (Rudel); koneill@aesop.rutgers.edu (O’Neill); mmcdermott@aesop.rutgers.edu (McDermott); Department of Agri-
Downloaded by [The University of British Columbia] at 21:15 03 February 2013

cultural, Food, and Resource Economics, Rutgers University, 55 Dudley Road, New Brunswick, NJ 08901, e-mail: gottlieb@aesop.rutgers.edu
(Gottlieb); Biological Sciences, State University of California–Chico, Chico, CA 95929, e-mail: chatfield@csuchico.edu (Hatfield).