ECOLOGY AND POPULATION BIOLOGY

Ecology of Urban Arthropods: A Review and a Call to Action

NANCY E. MCINTYRE
Center for Environmental Studies, Arizona State University, Box 873211, Tempe, AZ 85287Ð3211

Ann. Entomol. Soc. Am. 93(4): 825Ð835 (2000)

ABSTRACT A review of the entomological literature revealed relatively few general studies on
arthropods in urban environments, excluding those in the context of pest control or epidemiology,
and all were limited in scope and duration. Most studies documented the presence and abundance
of species in a variety of poorly quantiÞed urban categories. There also were a number of studies
on the effects of urban pollution and changes in arthropod community composition over time
(particularly in urban green areas). From these studies, three groups of arthropods could be
identiÞed: (1) “rural” taxa not present (or at lower abundance) in urban settings, (2) “urban” taxa
present only (or at higher abundance) in urban settings, and (3) taxa present in both rural and urban
settings with no particular afÞnity for either. The lack of a basic understanding of the mechanisms
accounting for distributional and abundance patterns of urban arthropods illustrates the many
opportunities for entomological research that exist in urban settings. Some of these opportunities are
outlined to encourage further work on the ecology of urban arthropods.

KEY WORDS urban ecology, urban entomology, urbanization

ARTHROPODS HAVE BEEN associated with humans and
their activities and constructions since recorded his-
tory (Curran 1937, Montgomery 1959, Hogue 1987,
Cloudsley-Thompson 1990). Despite this long associ-
ation, however, relatively little research has been
done on how arthropods use urban habitats. An urban
environment is a heterogeneous mosaic of residential
dwellings, commercial properties, parks, and other
land-use types that provides an array of habitat types
that can be used by arthropods. Although there have
been some studies done on arthropods in certain ur-
ban settings, such as landÞlls (Crawford 1979), resi-
dential yards (Owen 1971, Owen and Owen 1975), and
recreational parks (Faeth and Kane 1978, Sa-
woniewicz 1986, Kozlov 1996), few studies have com-
pared arthropods in various urban land-use types, and
those few have focused on speciÞc arthropod species
or families rather than on broader patterns of diversity
(Ehler and Frankie 1979a, b; Vincent and Frankie
1985). Although there is an enormous body of litera-
ture on synanthropic arthropod species that transmit
diseases to humans or livestock or that harm agricul-
tural crops (see reviews in Olkowski et al. 1976, Ebel-
ing 1978, Frankie and Ehler 1978, and Dreistadt et al.
1990), there has been surprisingly little attention paid
to how urban development affects the abundance and
diversity of nonpest arthropods and of arthropods in
general. This lack of attention to urban environments
in entomological research reßects a more widespread
pattern of overlooking urban environments in the nat-
ural sciences in general (Matson 1990, McDonnell
1997, Walbridge 1997). Urbanization is considered
one of the primary causes for declines in arthropod
populations, however, making the study of arthropods
in urban environments an extremely important and
timely avenue for research (Pyle et al. 1981).
Effects of Urbanization on Arthropods
Urbanization is associated with a variety of effects
on arthropods, including pollution, drainage and di-
version of watercourses, and habitat fragmentation
and loss (see discussion in Pyle et al. [1981]). These
factors have led to declines and even extinctions of
some arthropods (e.g., the loss of coastal sand dunes
to urban development around San Francisco, CA,
USA, has been implicated in the extinction of three
butterßy species [Tilden 1956]). Urbanization also
usually involves the introduction of exotic ßora for
landscaping. Some exotic arthropods have been inad-
vertently introduced in landscaping materials (Nowa-
kowski 1986, Barthell et al. 1998, Krell and Hangay
1998); these exotic species may then displace native
arthropods (Suarez et al. 1998). There may thus be
both direct and indirect effects from urbanization on
arthropods.
Why Study Arthropods in Urban Environments?
Arthropods are particularly logical choices for
studying effects from urbanization for the following
Þve reasons: (1) they are diverse and thus may provide
a snapshot of overall biological diversity in an area; (2)
their relatively short generation times mean that they
may respond quickly to anthropogenic changes to soil
and vegetation; (3) they are fairly easy to sample (and
many of the general public do not object to sampling
arthropods as opposed to vertebrates); (4) they rep-
resent a spectrum of trophic levels; and (5) they may

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826 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 93, no. 4

be important sociological, agronomical, and econom-

ical components of human-altered habitats. Any
changes in their environment may disrupt the func-
tions that arthropods play in the ecosystem. By the
year 2000, an estimated 50% of the worldÕs human
population is expected to live in urban settings (Rod-
ick 1995); and with the worldÕs annual human popu-
lation growth rate at 1.8% (which translates into ⬇90
million more people each year [Matson 1990]), urban
development will accelerate in the foreseeable future.
How will arthropods be affected by this trend? Unless
work begins now on arthropods in urban environ-
ments, we may miss the opportunity to answer this
question.

Objectives
Although there have been previous reviews of ar-
thropods in urban environments (Frankie and Ehler
1978, Dreistadt et al. 1990), they focused mainly on
insects rather than terrestrial arthropods as a whole,
and both considered insects primarily with respect to
pest control and epidemiology in urban environments.
There are also several edited volumes that can be
considered reviews, although they, too, are dominated
by papers dealing with management of pest species
(e.g., Frankie and Koehler 1978, 1989; Bornkamm et al.
1982). Another review by Zapparoli (1997) did con-
sider the general ecology of insects in urban environ-
ments, but this review was conÞned to the insects of
a portion of Rome, Italy. In this current review I
summarize the ecological research done on urban
arthropods, other than in respect to pest control and
epidemiology, to determine any general patterns of
how arthropods respond to urbanization. This focus is
appreciably different from those taken in previous
assessments of arthropods in urban environments. I
will also discuss some of the many opportunities that
exist for future work on arthropods in urban environ-
ments. My primary objective is to generate impetus for
studies on the ecology of urban arthropods.

Types of Papers Reviewed

In my literature search I found 79 peer-reviewed
journal articles on the ecology of arthropods in urban
environments, dating from 1933 to 1999. I excluded
papers that considered urban arthropods solely as ep-
idemiological vectors and papers evaluating the efÞ-
cacy of pest-management methods. This criterion did
not exclude papers that included pest species if the
primary focus of the paper was not pest control per se
(e.g., Hanks and Denno 1993, 1994). All the papers
used included at least an abstract in English. Although
there probably is a fair amount of literature about
urban arthropods written in languages other than En-
glish, this literature is unfortunately poorly known.
The papers that were reviewed represent studies from
⬎51 cities in 14 countries on six continents. For each
paper, I identiÞed the urban land-use type(s) present
and the arthropod taxa studied. I summarized the
conclusions from each paper and tallied any similar-
Fig. 1. Proportion of studies devoted to particular ar-
thropod taxa.
ities among papers. I also looked for trends in study
topics or methodology over the years.
Results of Review
Scope and Duration of Studies. For most cases,
there was insufÞcient replication to tally results by site
or taxon to determine if any population and commu-
nity responses to urbanization are repeated across
different cities, countries, or continents. Most studies
focused on Lepidoptera and Coleoptera, and most
arthropod orders were not included (Fig. 1). In ad-
dition, many studies did not identify arthropods be-
yond the order or family level. These shortcomings
illustrate the need for more work on arthropods in
July 2000 MCINTYRE: URBAN ARTHROPODS
Fig. 2. Number of studies conducted in various types of
land use. Some studies were conducted in more than one type
of land use, therefore the total frequency is greater than the
number of studies reviewed (79).
urban environments in general. In addition, most au-
thors did not include raw data in their papers upon
which to perform statistical meta-analyses. All studies
reviewed were rather limited in scope and duration.
Only 11 studies were conducted in more than one
urban area, and no study lasted longer than 3 yr (and
most had only one sampling period). In addition, most
of the studies were observational, not experimental.
Although such studies provide useful information,
long-term manipulative studies are needed to deter-
mine the mechanisms accounting for why populations
and communities of urban arthropods differ from rural
ones.
Study Site Quantification. Most studies were con-
ducted in forests or other green areas; relatively few
studies were conducted in more built-up areas (Fig.
2). Before 1985, all of the papers presented urbaniza-
tion qualitatively; that is, the authors made compari-
sons among categories of urbanization, such as “park”
and “suburb.” Some of these studies considered sites to
exist along a gradient of variables such as pollution or
human population density (Bishop et al. 1975,
Czechowski 1982, Klausnitzer and Richter 1983,
Ruszczyk 1986, Sustek 1987, Ruszczyk and de Araujo
1992, Pouyat et al. 1994, Blair and Launer 1997, Denys
and Schmidt 1998), although actual quantiÞcation in
gradient or site descriptions (e.g., measuring percent
vegetation cover, percent pavement) was present in
papers only after 1985 (Ruszczyk 1986, Ruszczyk and
de Araujo 1992, Pouyat et al. 1994, Schmitz 1996, Blair
and Launer 1997, Denys and Schmidt 1998). Even so,
not all papers after 1985 quantiÞed their study sites.
Although some qualitative descriptions may be highly
detailed (e.g., Czechowski 1982), more rigorous site
descriptions will be needed to determine the mech-
anisms accounting for patterns of synanthropization
and also to ensure proper replication and repeatability
of studies over time. QuantiÞcation is particularly im-
827
portant when considering that different areas of open
space may not be equally attractive to arthropods, and
these differences would only be detected if aspects of
“open space” are quantiÞed. For example, certain
grasshoppers (Orthoptera: Acrididae) and mining
bees (Hymenoptera: Andrenidae) are more common
in parks than in residential yards, even though both
areas are often considered collectively as “green areas”
(Gilbert 1989). Despite their superÞcial similarities,
such areas may differ in characteristics as grass height
and plant species diversity, which may differentially
attract or repel certain arthropods. Thus, it is impor-
tant to describe and quantify urban habitats in detail.
Features that should be measured in an urban study
site include the type and amount of vegetation, soil,
and built structures present; the height of vegetation
and buildings; presence of host plants and oviposition
sites; canopy cover; wind speed; humidity; water cur-
rent speed, pH, turbidity, oxygenation, and salinity;
human population density; and other environmental
characteristics (see reviews in Rowntree 1984, 1986;
Sutherland 1996). Statistical ordination techniques
such as correspondence analysis may be useful in
relating patterns of arthropod abundance and diver-
sity with quantiÞed urban habitat measurements, such
as percent ground cover or vegetation height (ter
Braak 1995). There is no standard methodology for
studies of arthropods in urban environments, nor
should there be given the variety of organisms, loca-
tions, and research questions. However, measurement
of study site variables will permit a deeper and more
precise understanding of the biological patterns and
mechanisms present.
Types of Studies on Urban Arthropods
A number of the studies were concerned with doc-
umenting speciesÕ presence and abundance, some-
times comparing different sites (e.g., Bristowe 1939;
Edelsten 1940; Woodroffe 1955; Klausnitzer and Rich-
ter 1983; Lazenby 1983, 1988; Kirby 1984; Sustek 1992,
1993). Most of these inventories were conducted in
“green areas” such as parks, residential yards, wood-
lots, and vacant lots and thus are probably not repre-
sentative of a heterogeneous urban environment as a
whole. For the studies that did not focus on composing
species lists, two general research themes emerged:
(1) effects of urban pollution on arthropods, and (2)
changes in arthropod communities over time during
vegetational succession.
Effects of Urban Pollution on Arthropods
Industrial Melanism and Air Pollution. Many of the
studies of the effects of urban pollution on arthropods
were concerned with industrial melanism (Owen
1962, Clarke and Sheppard 1966, Leblanc and
DeSloover 1970, Gilbert 1971, Bishop 1972, Muller
1972, Bishop et al. 1975, West 1977, Wiackowski 1978,
Creed et al. 1980, Clarke et al. 1985, Berry 1990, Stew-
art and Lees 1996, Cook et al. 1999). Industrial mel-
anism has been particularly well documented in the
828 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
British peppered moth (Lepidoptera: Geometridae,
Biston betularia L.), whereby increases in airborne
sulfur dioxide killed light-colored lichens on tree
trunks where the moths rested during the day, making
the moths more conspicuous against the dark trunks to
predators such as birds. In addition, tree trunks them-
selves became darkened from build-up of soot depos-
its. Through natural selection, light-colored moths de-
creased in frequency, whereas a darker-colored
morph became more abundant in woods near indus-
trial centers. Light-colored moths were still more
abundant in rural areas exposed to less air pollution
(Kettlewell 1955a, 1955b 1956, 1958). More rigorous
industrial emissions standards have decreased the
amount of airborne SO2, which has permitted re-
growth of lichens and a shift in selection to favor
light-colored morphs over darker ones once again in
many areas of Britain and the United States (Grant et
al. 1996, 1998). Although certain authors (e.g., Sargent
1968, Clarke et al. 1994, Coyne 1998, Sargent et al.
1998) have found fault with various aspects of this
system, the existence of industrial melanism is indis-
putable (see reviews in Kettlewell 1973, Majerus 1998,
Grant 1999).
The pollution-related decline of tree lichens has
caused declines in other arthropods from trophic ef-
fects. For example, lichenophages and herbivores (es-
pecially Psocoptera) decreased in British woods that
experienced a pollution-related lichen die-off, al-
though some omnivores (Isoptera, Dermaptera, and
Diptera: Psychodidae) actually increased in abun-
dance (Leblanc and DeSloover 1970, Gilbert 1971,
Wiackowski 1978). Industrial melanism was also once
thought to account for the dominance of dark-colored
morphs of the two-spot ladybird beetle (Coleoptera:
Coccinellidae, Adalia bipunctata L.) in urban areas
compared with more rural sites (Creed 1971, 1974),
although it is now believed that a dark morph in this
species is actually an adaptation to capture solar heat
efÞciently in areas that may be cooled from smoke
clouds (see review in Muggleton et al. [1975]). Air
pollution (particularly from sulfurous compounds)
has been implicated in declines in spiders in London,
England (Bristowe 1939), mites and springtails in
Switzerland (Steiner 1995), and overall insect diver-
sity in Poland (Wiackowski 1978). A decline of ßying
insects in London that led to concomitant declines in
insectivorous birds was attributed to air pollution;
improvements in air quality are thought to have led to
an increase in both insects and birds (Cramp and
Tomlins 1966, Cramp and Gooders 1967, Gooders
1968). Although heavy metals from vehicle exhaust
have not been found to decrease soil and litter faunal
diversity in roadside verges (Williamson and Evans
1973), heavy metals from other sources have been
linked to declines in arthropod diversity elsewhere
(see review of effects of heavy metals on biodiversity
in Tyler et al. [1989] and Pouyat et al. [1994]).
Thermal Pollution and Latitudinal Gradients. In a
study by Tischler (1973), the effect of urban “thermal
pollution” (i.e., cities as “heat islands” from both in-
creased production of heat from concentration of peo-
Vol. 93, no. 4
ple and machines as well as increased heat reßectance
and absorbency from pavement [Kim 1992]) offset a
latitudinal gradient in the population density and cy-
cling of various arthropods in Europe, so that popu-
lations in cities at more northern latitudes were more
similar to those in rural settings at more southern
latitudes than rural populations farther north. A sim-
ilar example of how urbanization can counteract lat-
itude has been noted for bird diversity (Jokimaki and
Suhonen 1993).
Host-Plant Stress from Pollution and Human Ac-
tivity. The presence of pollution (particulate or ther-
mal) means that the biota in and around urban areas
are subjected to different stresses than would be the
case in more isolated areas. For example, because host
plants in urban settings are exposed to different am-
bient conditions than are the same plant species in
more rural areas, host plants may differ in their “at-
tractiveness” to herbivorous insects between these
two settings. Insect diversity and densities on those
plants may consequently differ between urban and
rural areas. This appears to be the case for a species of
scale insect (Homoptera: Coccidae, Pulvinaria regalis
Canard). Soil compaction and the presence of con-
crete made the ground surface impenetrable to pre-
cipitation and nutrients, which affected the health of
the scaleÕs host trees in urban areas and, thus, in-
creased their susceptibility to scale colonization
(Speight et al. 1998). A similar kind of relationship was
discussed by Schmitz (1996), who speculated that
aster plants (Asteraceae) that are stressed from lack of
water create sap with higher concentrations of free
amino acids than do watered plants. Urban asters may
receive less water than rural plants because of soil
compaction and distance from natural water sources
such as rivers or swamps. Thus, concentrated sap from
urban plants may be a higher-quality resource to
plant-sucking insects such as aphids (Homoptera:
Aphididae), leading to higher aphid densities on urban
than on rural plants (Schmitz 1996; but see Connor
1988, Hanks and Denno 1993, Nuckols and Connor
1995). Additionally, urban plantings that are scattered
may be unable to support high densities of predatory
arthropods (because of dispersion of prey), enabling
urban herbivores (e.g., scale insects, Homoptera: Di-
aspididae) to attain higher densities than in natural
habitats (Hanks and Denno 1993). In another exam-
ple, holly leafminers (Diptera: Agromyzidae) were
more abundant on urban than rural holly trees near
Newark, DE (USA), because the urban trees were
exposed to more sunlight (not being shaded by over-
story trees, as they would be in a natural forest set-
ting), which hastened leaf senescence and abscission.
Although this process did not affect the leafminers per
se (which completed their larval development in the
fallen leaves), it did lower rates of parasitism from
hymenopteran parasitoids, which did not search for
hosts among fallen leaves. Lower rates of parasitism
among the urban leafminers thus promoted higher
leafminer densities in urban areas than in rural forests
(Kahn and Cornell 1989). Regular pruning of plants in
urban settings may stimulate new plant growth,
July 2000 MCINTYRE: URBAN ARTHROPODS
thereby providing consistent tender forage for cater-
pillars (Owen 1971). These studies illustrate how ur-
ban areas may actually support higher numbers of
phytophagous insects than would otherwise be ex-
pected in such an altered environment, but more re-
search is needed to determine whether this is a general
pattern and to rule out the importance of other mech-
anisms (e.g., urban plants may support higher densi-
ties of phytophages than do rural plants simply be-
cause there are fewer plants upon which to settle in
urban areas).
Water Pollution. Just as air and soil characteristics
may differ between urban and rural areas, water qual-
ity may similarly be affected by pollution and drainage
or diversion of watercourses. These habitat changes
may thus affect aquatic arthropods. For example,
benthic arthropod communities differed among
streams that ran through settlements with different
human population densities (Jones and Clark 1987).
Although upstream human population density was not
a good predictor of the total number of arthropods or
arthropod density, urban streams were dominated by
midges (Diptera: Chironomidae), whereas more rural
streams had a more diverse arthropod community.
Streams near larger human settlements differed from
streams near fewer people in terms of several abiotic
variables (such as dissolved oxygen, alkalinity, sedi-
ment load, heavy-metal concentration), and midges
appear to be more tolerant of the urban conditions
(Jones and Clark 1987). Similarly, fewer arthropods
were found near human-frequented boardwalks in
mangrove swamps around Sydney, Australia, than in
more secluded areas farther away from the board-
walks, although the reasons for this pattern are not
fully understood (Kelaher et al. 1998). Finally, Claas-
sen (1933) discusses how pollution from such sources
as milk treatment factories and salt puriÞcation cen-
ters in New York perturb stream arthropod commu-
nities to the detriment of the entire stream ecosystem.
Changes in Arthropod Communities Over Time
During Vegetational Succession
Habitat alterations that result from urbanization
affect the arthropods present. As urbanization con-
tinues over time, there are concomitant changes in
arthropod community composition over time. Sites
that are cleared and left undeveloped attract different
species as they undergo vegetational succession. For
example, the arthropod communities in 12 vacant lots
of different ages were compared in ShefÞeld, England
(Gilbert 1989). Overall diversity declined from the
oldest sites (12-15 yr since last disturbance), to inter-
mediate-aged sites (4 Ð 6 yr), to the most recently
cleared sites (0 Ð1 yr). In general, recently cleared
sites are Þrst dominated by generalists and predators
(especially Coleoptera: Carabidae, Staphylinidae; and
Araneae), particularly species that are vagile and thus
good dispersers (Lazenby 1988, Gilbert 1989). Sites in
early successional stages often are dominated by pred-
ators (Odum 1969). The presence of certain species in
early successional sites may be the result of proximity
829
to nearby undisturbed sites that are sources of colo-
nists, such as Lepidoptera (particularly Hesperiidae
and Lycaenidae), Hymenoptera (Formicidae, Bom-
bidae), Coleoptera (Carabidae), Araneae (Linyphi-
idae), and Isoptera (Gilbert 1989). These pioneers
come from surrounding areas, particularly other areas
of open space within a 1-km radius (Davis 1979). Over
time, more sessile, specialist species can be supported,
depending on the availability of host plants and other
resources (Brown 1985, Vincent and Frankie 1985; cf.,
Denys and Schmidt 1998); arthropod communities of
later seral stages are usually more even in terms of
species abundance (Horn 1974), and body sizes (Sus-
tek 1987, 1993). As succession proceeds, carabids de-
cline in abundance and other beetle families (partic-
ularly Buprestidae, Chrysomelidae, Coccinellidae,
and Curculionidae) increase in abundance (Lazenby
1988). Similarly, linyphiid spiders decline after 4 Ð15 yr
as Thomisidae, Lycosidae, and Araneidae become
more abundant (Gilbert 1989). Exotic “urban special-
ist” arthropods may become extremely abundant with
time, such as the cockroach Blatta orientalis L. (Blat-
taria: Blattidae) and the theridiid spider Enoplognatha
thoracia (Hahn) (Araneae: Theridiidae) (Bateson and
Dripps 1972, Crawford 1979). Total urban arthropod
density may reach as high as 3,854 individuals per
square meter even without the presence of exotic
invaders (Majzlan and Holecova 1993).
Although many references state that some taxa “de-
cline in abundance” with seral age, a distinction should
be made between numeric and proportionate de-
clines. In a numeric decline, a taxon decreases in
absolute number. In a proportionate decline, a taxon
decreases in its importance to an overall assessment of
diversity. In such a case, a taxon may be increasing in
absolute number but be only one of an increasing
number of taxa present. In some cases (e.g., Gilbert
1989), a numeric decline is used; in other cases (e.g.,
Lazenby 1988), a proportionate decline is meant but
it is unclear whether a numeric decline is also present.
Therefore, it is important to note the distinction.
Arthropods provide a good system for studying suc-
cessional processes within urban patches because just
as patches may undergo vegetative succession within
an urban setting, urban development can itself be
considered a form of succession. Changes in land-use
may be frequent in urban areas because parcels of land
are constantly being annexed, zoned, and built upon
or razed and built upon again. Urbanization has tra-
ditionally been considered a form of environmental
disturbance (Rebele 1994). Several of the studies re-
viewed here were conducted in parcels of land sur-
rounded by urban development but not themselves
developed (i.e., “open space” or “green areas”). Island
biogeography theory, which explains how diversity is
related to the area and isolation of a site (be it an
oceanic island or an “island” of open space surrounded
by buildings [MacArthur and Wilson 1967]), was the
research paradigm for most of these studies (e.g.,
Faeth and Kane 1978, Kozlov 1996). In some cases, the
arthropod community was described as being similar
in composition among sites (Ehler and Frankie 1979a,
830 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
1979b; Dawe and McGlashan 1987), at least for those
sites similar in area (Faeth and Kane 1978, Davis 1979,
Nowakowski 1986). Most of these studies emphasized
that green areas promote diversity by providing areas
free from disturbance (Okuma and Kitazawa 1982,
McGeoch and Chown 1997). Other studies, however,
noted that certain species seem to prefer sites kept at
an early successional stage through such activities as
periodic mowing (Tischler 1973, Czechowski 1982); in
these cases, urbanization was considered to manifest
itself as a chronic disturbance. One study noted that
species richness peaked at intermediate levels of ur-
banization/disturbance, as per the “intermediate dis-
turbance hypothesis” of Connell (1978) (Blair and
Launer 1997), and another noted a similar effect in
that disturbance favored exotic ruderal species such as
the “pavement ant” (Hymenoptera: Formicidae, Tet-
ramorium caespitum L.) that would otherwise be com-
petitively excluded by native species (King and Green
1995).
Effects of Habitat Heterogeneity on Arthropod
Distributions in Urban Environments
Urbanization creates a heterogeneous mosaic of
patches consisting of built-up areas, areas of construc-
tion, and remnants of indigenous habitat. Some ar-
thropods may be able to exploit relatively small and
isolated habitat remnants, thereby being able to tol-
erate a greater degree of urban development than can
species that are more sensitive to habitat area and loss.
The exotic Argentine ant Linepithema (⫽Iridomyrmex)
humile (Mayr), for example, is able to exploit relatively
small patches of riparian and coastal scrub habitats near
highly developed areas and patches that are dominated
by exotic plants. The native ants, in contrast, appear to be
much more sensitive to habitat area and disturbance,
because they are found only in larger patches in more
rural areas with indigenous vegetation (Ward 1987, Su-
arez et al. 1998). A similar pattern has been found in
spider communities in urban woodlots in Japan, where
smaller forests had fewer species than did larger forests,
possibly because of the lack of larger prey items in the
smaller forests (Miyashita et al. 1998). In contrast, many
carabid beetle species were found at higher densities in
smaller forest fragments than in contiguous forests in
Finland, probably because the smaller habitat remnants
possessed higher vegetational diversity caused by re-
peated human disturbance and successional changes
(Halme and Niemela 1993). Most of the carabids cap-
tured in the smaller fragments were habitat and dietary
generalists that also occurred in a variety of other hab-
itats, whereas the species from the contiguous forests
were specialists (Halme and Niemela 1993).
Some studies noted that arthropods may be distrib-
uted patchily within a city, reßecting heterogeneity of
urban land-use types (e.g., Czechowski 1982, Kozlov
1996). Some authors attributed high arthropod diver-
sity to the presence of high vegetative and structural
diversity (Owen 1971, Clark and Samways 1997),
which follows from theories of MacArthur and
Vol. 93, no. 4
MacArthur (1961). This relationship may actually re-
ßect the nearby availability of resources such as host-
plants, however, rather than vegetative diversity per
se. Although grass lawns supported high densities of
Collembola (Natuhara et al. 1994), even relatively thin
strips of lawn isolated populations of more sessile taxa
(Clark and Samways 1997). Absence of leaf litter in
urban areas was thought to expose overwintering ar-
thropods to cold weather; presence of leaf litter was
associated with higher diversity of most microarthro-
pods (Davis 1978, 1979).
Although some studies noted no changes in diver-
sity or density with proximity to a city (Lussenhop
1973, Burakowski and Nowakowski 1981, Pouyat et al.
1994), other studies noted that different numbers of
species were encountered at different distances from
the city center. For example, Davis (1978, 1979) doc-
umented an ⬇60% decline in species richness with
proximity to the center of London, England. Declines
in species richness with increasing urbanization were
attributed to several factors, including isolation from
colonizing source populations from natural habitats
(Davis 1979, Denys and Schmidt 1998), pollution
(Pouyat et al. 1994), and loss of foodplants (Ruszczyk
1986). In other studies, however, higher species rich-
ness was documented in centrally located sites than in
rural ones; this pattern was attributed to the presence
of specialist urban “invaders” (Kozlov 1996), species
accidentally introduced by humans from rural areas
(Nowakowski 1986), or the greater abundance of re-
sources in urban settings (Schmitz 1996). For exam-
ple, the number of species of phytophagous carabid
beetles was 2.5 times higher and overall abundance
was 20 times higher in green areas of Warsaw than in
a more rural forest in Poland; this pattern was attrib-
uted to the more consistent availability of foodplants
in the city throughout the year (Czechowski 1982).
Three Community-Level Responses to Urbanization
Based on species-speciÞc variation in arthropodsÕ
responses to urbanization, arthropods can be consid-
ered to belong to one of three distinct communities:
(1) rural taxa not present (or at signiÞcantly lower
abundance) in urban settings, (2) taxa present only
(or at higher abundance) in urban settings (i.e., synan-
thropic species), and (3) taxa present in both rural and
urban settings with no particular afÞnity for either
(see also Shorrocks 1969, Czechowski and Mikolajc-
zyk 1981, Blair and Launer 1997, Guarisco 1999). Rel-
atively few studies compared arthropod abundance
between urban and rural settings. Those that did
found that many taxa are found in both urban and rural
settings, but certain taxa are consistently more abun-
dant or conspicuous in one or the other setting (Table
1). For example, herbivores often were more abun-
dant in cities than in rural sites, particularly those
species that were specialists on horticultural plantings
or ruderal plants (Schmitz 1996, Denys and Schmidt
1998), whereas parasitoids usually exhibited the re-
verse pattern (Sawoniewicz 1986, Denys and Schmidt
July 2000 MCINTYRE: URBAN ARTHROPODS 831

Table 1. Examples of the three community-level responses Indicators of Urbanization

(rural, urban, or either) to urbanization
Presence and abundance of the members of these
Citation(s) Taxon
Community three communities can serve as “indicators” of differ-
type ent degrees of urbanization (Czechowski 1982).
Sawoniewicz 1986, Parasitoid Rural These indicators can be used to detect subtle prox-
Denys and Schmidt Hymenoptera imity effects of urbanization, even in sites that are not
1998 considered urban per se (such as sites on the fringe of
Kozlov 1996 Micropterix calthella Rural
L., Lepidoptera: developed areas). Indicator taxa should preferably be
Micropterigidae locally abundant and relatively easy to identify taxo-
Bateson and Dripps 1972 Blatta orientalis L., Urban nomically (Clark and Samways 1997); thus, local re-
Blattaria: Blattidae search is necessary to identify potential candidates
Crawford 1979 Enoplognatha Urban
thoracia (Hahn), that would be appropriate for a given region. How-
Arachnida: ever, some arthropod taxa were consistently catego-
Theridiidae rized as belonging to one of the three communities,
Ward 1987, Suarez et al. Linepithema humile Urban even in different regions of the world. For example,
1998 (Mayr),
Hymenoptera: carabid beetles (Coleoptera: Carabidae) were con-
Formicidae sidered to be indicators of urbanization in South Africa
King and Green 1995 Tetramorium Urban (Clark and Samways 1997), Poland (Czechowski
caespitum L., 1982), Germany (Klausnitzer and Richter 1983), Slo-
Hymenoptera:
Formicidae vakia (Sustek 1987, 1992), and Great Britain (Lazenby
Kozlov 1996 Caloptilia syringella Urban 1983). Carabids have often been described as “habitat
F. and Coleophora indicators” (Luff et al. 1992, Rykken et al. 1997).
sibiricella Flkv., Therefore, future studies on arthropods in urban en-
Lepidoptera:
Gracillariidae vironments may wish to examine closely how carabids
Ruszczyk 1992 Lepidoptera: Either respond to land-use changes. Other taxa (e.g., Or-
Papilionidae, thoptera: Gryllidae; Coleoptera: Cicindelidae; Diptera:
Pieridae Syrphidae and larvae from various other families; and
Kozlov 1996 Parornix scoticella Either
Stt., Lepidoptera: parasitic Hymenoptera) may also prove to be useful
Gracillariidae indicators of various effects of urbanization (see discus-
sion in Clark and Samways [1997]). The taxa denoted
“urban” or “rural” in Table 1 may also serve as indicators
(presence of “urban,” absence of “rural”).
1998). For the urban-dwellers, urban settings pro-
vided resources that were scarce in rural areas (Tis-
chler 1973). For example, rubbish and sewage pro- General Patterns of How Arthropods Respond to
vided food resources to many arthropods. In many Urbanization
cases, these resources offset negative effects of urban-
Arthropods respond both directly and indirectly to
ization, such as pollution (Czechowski 1982). Mem- urbanization. Arthropods respond directly to habitat
bers of this group were often considered preadapted loss and pollution (through direct mortality). Arthro-
to urban life because of morphological characteristics pods respond indirectly to urbanization in responding
such as high vagility (especially ability to ßy, which to alterations in habitat structure and in the presence
enabled them to disperse from natural sources to col- and abundance of resources (e.g., host plants). Both
onize urban areas) and small body size (which al- direct and indirect effects act on individuals (espe-
lowed them to take advantage of the small-scale re- cially individuals belonging to certain sensitive taxa),
sources characteristic of urban areas, such as which results in changes in local population density
individual plants in residential yards) (Tischler 1973, and community structure. Arthropod communities in
urban environments tend to be more diverse (owing
Ruszczyk 1986). Some urban species actually showed
to the presence of exotic taxa) than are communities
physical adaptations to urban life, such as ability to in nonurbanized areas, but not all portions of a city are
digest new larval host-plants (Owen 1971, Ruszczyk equally diverse because of differences in distances
1986), but more research is needed to determine from sources of arthropod colonists, land-use types,
whether speciÞc genetic adaptations have occurred in ages of development, and seral stages. Some taxa ben-
urban populations of arthropods (Tischler 1973, eÞt from urbanization, whereas others are negatively
Hanks and Denno 1994). Urban species also tended to affected or remain unaffected.
have large geographic ranges, indicating adaptability
to many regions and habitats (Shapiro and Shapiro
1973, Czechowska and Bielawski 1981). It is currently Future Directions of Research on Arthropods in
unknown how most arthropod taxa respond to urban- Urban Environments
ization (positively, negatively, or neutral), which is a Given these general patterns of how arthropods
topic that merits further investigation. respond to urbanization, it is now possible to formu-
832 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
late some testable hypotheses that merit investigation.
Some of these hypotheses include the following.
● Arthropod diversity should decrease as the level of
pollution (terrestrial, aquatic, or atmospheric) in-
creases.
● Habitat (e.g., a particular host-plant) that is scarce
and widely dispersed should experience higher rates
of occupation (herbivory, oviposition) than should
habitat that is abundant and widespread.
● Areas that have only recently been urbanized
should possess relatively more “early successional”
taxa (e.g., Lepidoptera: Hesperiidae, Lycaenidae;
Hymenoptera: Formicidae, Bombidae; Isoptera)
than should an older area. Predators (e.g., Araneae:
Linyphiidae; Coleoptera: Carabidae, Staphylinidae)
should be particularly well-represented in recently
developed sites.
● Diversity should increase with the age of an urban-
ized area.
● Urban sites that are nearer to undeveloped areas of
indigenous habitat should be more accessible to
colonizing arthropods than would be sites that are
more centrally located in an urban area; therefore,
areas on the fringe of development should possess
relatively higher diversity than more insular areas.
Much of the diversity in fringe areas will be repre-
sented by highly vagile taxa (the colonists), whereas
insular sites will have a more even representation of
mobilities and body sizes.
● The diversity of exotic species will increase with the
age of an urbanized area.
Urbanization is no fad, and entomology and ecology
should take advantage of the research opportunities
made available by the “natural experiment” (Diamond
1986) of urban development. Currently, there is a
paucity of work on synanthropic arthropods in South
America, Africa, Asia, and Australia, even though
these areas are experiencing some of the fastest rates
of human population growth and urban development.
Addressing the following questions in these and other
urban areas would be particularly useful.
● How do most arthropods respond to urbanization
(positively, negatively, or neutral)?
● How large do urban green areas need to be to pre-
serve rural levels of arthropod diversity?
● Do exotic arthropod species replace and Þll the
niches of native species? Are certain niches more
resilient or vulnerable to urban change than others?
How are these relationships mediated by land-use
type? How does the presence of exotic vegetation
mediate these relationships?
● Are native arthropods able to maintain their popu-
lation levels without supplementation from rural
source populations? Are population dynamics dif-
ferent for species near the urban fringe than those
near the urban center? How does the proximity of
remnants of undeveloped open space affect these
relationships?
● How do socioeconomic variables interact with phys-
Vol. 93, no. 4
ical (habitat) variables to inßuence arthropod di-
versity and abundance?
Some of these questions are being addressed by a
new long-term research project monitoring arthropod
diversity and turnover in various types of urban land
use in Phoenix, AZ, USA (http://caplter.asu.edu/re-
search/projects).
Conclusions
The difÞculties that will surely be encountered in
the identiÞcation and study of many synanthropic
arthropods do not diminish the importance of under-
standing arthropods in urban environments. Given
that arthropods are being affected by urbanization at
an unprecedented rate, the important roles they play
in ecosystem structure, function, and dynamics and in
comprising the majority of the EarthÕs biodiversity are
also being altered at an unprecedented rate. Without
further research on how arthropods use urban envi-
ronments, however, we cannot predict how these
functions may change with future patterns of urban
development. Without such knowledge, we will be
unable to plan urban development to facilitate arthro-
pod conservation, a subject whose importance is in-
creasingly being recognized as humans continue to
modify the EarthÕs ecosystems (Samways 1992).
Acknowledgments
Comments from S. Faeth, W. Fagan, and J. Rango (De-
partment of Biology, Arizona State University) and an anon-
ymous reviewer improved earlier versions of the manuscript.
This paper is a product of the National Science FoundationÕs
Central Arizona-Phoenix Long-term Ecological Research
Project (DEB-9714833).
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