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Urban wildlife research: Past, present, and

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Article in Biological Conservation October 2012

DOI: 10.1016/j.biocon.2012.06.018

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 Biological Conservation 155 (2012) 2332

Contents lists available at SciVerse ScienceDirect

Biological Conservation
journal homepage: www.elsevier.com/locate/biocon

Review

Urban wildlife research: Past, present, and future

Seth B. Magle a,, Victoria M. Hunt b, Marian Vernon a, Kevin R. Crooks c
a
Lincoln Park Zoo, Urban Wildlife Institute, Department of Conservation and Science, Chicago, IL 60614, United States
b
Department of Biological Sciences, University of Illinois at Chicago, Chicago, IL 60607, United States
c
Department of Fish, Wildlife, and Conservation Biology, Colorado State University, Fort Collins, CO 80523, United States

a r t i c l e i n f o a b s t r a c t

Article history: Despite expanding urban areas and increased awareness of urbanization impacts on wildlife, trends in
Received 14 March 2012 urban wildlife studies have not been evaluated systematically. We performed a thorough assessment
Received in revised form 5 June 2012 of such research, evaluating urban wildlife publications from 16 leading journals in animal behavior, con-
Accepted 12 June 2012
servation, ecology, general science, landscape ecology, and wildlife biology from 1971 to 2010. Using a
Available online 17 July 2012
systematic review process, we quantied trends in urban wildlife research over time and in different sci-
entic elds, and also assessed author afliations, geographic and taxonomic focus, research topics, and
Keywords:
study site types. In general, rates of publication for urban wildlife research have been increasing,
Urban
Wildlife
although still remain low (<2% of publication volume) considering urban growth and its impacts on bio-
Literature review diversity are accelerating globally. Landscape ecology and wildlife biology journals, followed by conser-
Temporal trends vation journals, published the highest percentage of urban wildlife publications, whereas such studies
Geographic were rare in animal behavior, ecology, and general science journals. Academics were rst-authors on
Publication ca. 75% of urban wildlife publications, whereas research directed by government agencies, non-govern-
mental organizations, and private industry were less common, with little evidence of temporal shifts
in these patterns. The majority of urban research studied birds or mammals, and nearly all was conducted
in North America, Europe, or Australia, as expected given expansive urban development and associated
research focus. Animal behavior was the most common scientic topic in urban wildlife research, fol-
lowed by conservation, landscape ecology, wildlife management, and population ecology. While subur-
ban and exurban development have been recently identied as an important issue, we found no
evidence that research in these study systems has increased. Author afliation, geographic location, tax-
onomic focus, and research topics of urban wildlife studies were generally similar to those conducted in
non-urban systems, although avian studies were more common, and African and community ecology
studies less common, in urban compared to non-urban areas. We suggest that the most critical gaps
for urban wildlife researchers are in rapidly urbanizing areas in South America, Africa, and Asia, and
on understudied taxa such as herpetiles, sh, and arthropods. Research conducted on multiple taxa
and across continents is also rare, but will be necessary for global understanding of ecological dynamics
of urban systems.
2012 Elsevier Ltd. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
2. Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
2.1. Journal selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
2.2. Publication inclusion/exclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
2.3. Publication categorization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
2.4. Non-urban publications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
2.5. Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
3. Results. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
3.1. Annual trends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
3.2. Journals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

Corresponding author. Tel.: +1 312 742 7215; fax: +1 312 742 7220.
E-mail address: SMagle@lpzoo.org (S.B. Magle).

0006-3207/$ - see front matter 2012 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.biocon.2012.06.018
24 S.B. Magle et al. / Biological Conservation 155 (2012) 2332

3.3. Author affiliation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

3.4. Geographic focus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
3.5. Taxonomic focus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
3.6. Scientific topic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
3.7. Research site types . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
3.8. Comparing urban and non-urban wildlife publications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
4. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
5. Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

1. Introduction also tend to be homogenized in cities, though overall diversity

The rate at which the world is urbanized is accelerating (Dear-
born and Kark, 2010; Forman, 2008; McKinney, 2002; Pickett et al.,
2008; Ramalho and Hobbs, 2012), and perhaps as a result, interest
in wildlife in urban areas appears to be growing (Adams, 2005; For-
man, 2008; Mayer, 2010; Mcdonald et al., 2008; Miller and Hobbs,
2002). Despite some study of urban plant species and a few iso-
lated investigations of urban wildlife in Europe, systematic re-
search on animal species in and around cities did not begin in
earnest until the 1970s (Gehrt, 2010; Sukopp, 2002). Since that
time, urban ecology has been formalized as an international, inter-
disciplinary eld (e.g., Marzluff et al., 2008), with several journals
devoted solely to the topic, and research boosted by several
high-prole government projects such as the urban Long-Term
Ecological Research sites in the United States (Grimm et al.,
2000), the urban group of the United Nations Educational, Scien-
tic, and Cultural Organizations Man and the Biosphere program
(Adams, 2005), and the Australian Research Centre for Urban Ecol-
ogy (ARCUE, Australian Research Centre for Urban Ecology, 2011).
Despite this emergent eld, however, relatively little is known
about historic and recent trends in urban wildlife research.
Urban ecosystems are not only increasingly common, but they
can play important ecological roles (Dearborn and Kark, 2010). This
may include protection of local biodiversity and rare or imperiled
species (Bolger et al., 1997; Crooks et al., 2004), either directly
for organisms that reside in urban areas or via creation of corridors
or stepping stone patches for individuals moving through cities
(Gibb and Hochuli, 2002; FitzGibbon et al., 2007). Moreover, in
addition to their role in ecology and conservation, urban green
space provides important benets to humans. These benets in-
clude ecosystem services such as pollination (Mendes et al.,
2008), air quality improvement (Nowak et al., 2006), and carbon
sequestration (Pickett et al., 2008), but also extend to simple
enhancement of property value (Bolitzer and Netusil, 2000;
Waddell and Moore, 2008). Perhaps one of the most powerful roles
of urban ecosystems is their ability to connect urban citizens with
their environment. It is often difcult for residents to develop an
appreciation for wildlife when conservation messages and pro-
grams focus on distant species or places (Orr, 1993). Urban habitats
allow people to engage directly with nature where they live and
recreate, and this approach may be particularly effective for nature
education and awareness (Chawla, 1999; Dearborn and Kark,
2010). The rate at which natural habitats are conserved for wildlife
is showing signs of decline (Blair and Johnson, 2008), and conse-
quently urban conservation areas will assume greater importance
in the future.
Urban areas are also characterized by their own unique chal-
lenges. Fragmented urban landscapes isolate natural populations
and restrict dispersal and functional connectivity, which can lead
to demographic and genetic instabilities (Frankham, 2006; Magle
et al., 2009) and community-level impacts (Crooks and Soul,
1999; Faeth et al., 2005; Crooks et al., 2010). Biotic communities
and abundance are often quite high (Chace and Walsh, 2006;
McKinney, 2006). Urban biota increasingly is comprised of non-na-
tive species that are both directly imported and benetted by ur-
ban land modication (McKinney, 2006; Simberloff, 1997).
Animals that reside in cities frequently conict with humans,
including incidents of animal-vehicle collision (Forman et al.,
2003), animal attacks on pets (Curtis and Hadidian, 2010; Gehrt
and Riley, 2010), and damage to lawns and landscaping (Urbanek
et al., 2011). In addition, urban areas, characterized by high densi-
ties of both humans and wildlife, represent a signicant risk for the
spread of disease (Bradley and Altizer, 2007). Zoonotic pathogens,
diseases that can spread from wildlife to humans, now constitute
75% of all emerging human diseases (Taylor et al., 2001).
To date, no review has provided a quantitative and comprehen-
sive evaluation of long-term trends in urban wildlife research
across a diversity of scientic disciplines and their associated jour-
nals. Herein, we adopt a systematic review process to evaluate
trends in urban wildlife research over time and in different scien-
tic elds, and also assess geographic (Chace and Walsh, 2006) and
taxonomic (Lawler et al., 2006) focus, including identifying gaps
where urban wildlife research is scarce. Given recent attention
on urbanization and its impacts, we expected a rapidly increasing
number of scientic publications focused on urban wildlife. We
also expected that birds would represent the majority of urban
wildlife studies (following Adams, 2005; Marzluff et al., 2001),
and that such research would be biased towards North America
and Europe given extensive urban growth and an associated focus
on urban research. We also investigated author afliation, predict-
ing that while academics conduct the majority of urban wildlife
research, studies directed by governmental agencies and non-gov-
ernmental organizations may be increasing as this is a stated goal
in urban ecology research (Young and Wolf, 2006). Further, we
evaluated studies conducted across a gradient of residential land
use types, including urban, suburban, and exurban systems,
expecting an increase in suburban and exurban studies given
heightened interest in development impacts outside of high-den-
sity urban areas (DeStefano and DeGraff, 2003; Theobald, 2001,
2005). Finally, we examined major research topics in urban wildlife
studies to identify primary scientic questions addressed by such
projects, and also compared such studies to those conducted in
non-urban systems. Our study reveals potential gaps in current
work, illustrates areas of strength and weakness, and provides
guidance for future urban wildlife research efforts.
2. Materials and methods
Our methods followed recommendations of Roberts et al.
(2006) for systematic reviews, although as we were assessing
trends as opposed to the effect of a process, policy, or intervention,
some components of the systematic review process were not appli-
cable. Specically, as we were interested in study focus rather than
 S.B. Magle et al. / Biological Conservation 155 (2012) 2332
outcomes, determining a hierarchy of evidence by which to assign
weight to publications was unnecessary.
2.1. Journal selection
First, we identied up to three target journals within the animal
behavior, conservation, ecology, landscape ecology, and wildlife
biology elds. We used Journal Citation Reports within the Insti-
tute for Scientic Information (ISI) Web of Science (Thomson
Reuters, New York, NY, USA) to rank journals based on impact
factor in 2000, using ISIs standardized subject categories of behav-
ioral science (for behavior journals), biodiversity conservation (for
conservation journals), and ecology (for ecology, landscape ecol-
ogy, and wildlife biology journals). We chose the year 2000 as it
represented an approximate midpoint in the period where we
anticipated high rates of urban wildlife publication (1990-present).
Within the ecology topic, we dened a wildlife biology journal as a
journal with wildlife in the title, and a landscape ecology journal
as a journal with landscape in the title. We omitted journals that
did not publish at least 50 articles in 2000 and those that did not
exist prior to 1990 to ensure adequate sample size for analysis,
and also omitted any journal that, based on its title, could not rea-
sonably be expected to regularly and frequently publish on wildlife
(e.g., Molecular Ecology, Behavioral and Brain Sciences). We selected
up to three journals from those that remained in each eld based
on the highest impact factor ratings; we selected the highest-tier
journals because they have the highest readership, and thus best
reect exposure of the scientic community to the topic. Following
these criteria, the journals selected from each ISI subject category
were: Animal Behaviour, Behavioral Ecology, and Behavioral Ecology
& Sociobiology for animal behavior; American Naturalist, Biological
Conservation, and Conservation Biology for biodiversity conserva-
tion; Ecology, Ecology Letters, and Journal of Applied Ecology for ecol-
ogy; and Journal of Wildlife Management, Wildlife Research, and
Wildlife Society Bulletin for wildlife biology. In the landscape ecol-
ogy eld, only two journals met our criteria, Landscape Ecology
and Landscape and Urban Planning. For comparative purposes, we
also included the two highest-impact general science journals, Sci-
ence and Nature, within the multidisciplinary science category in
ISI.
2.2. Publication inclusion/exclusion
Within each target journal, we performed a literature search
during AugustNovember 2011 within Web of Science, searching
for the term urban for all publications from 1971 to 2010. The
 notation ensured that terms such as urbanization, subur-
ban, exurban and semi-urban were also detected. Each publi-
cation that resulted from that search was then reviewed to
determine if it met the criteria for inclusion. Publications were in-
cluded only if they represented original research on wildlife (non-
domestic animals) located in human-dominated, non-agricultural
areas. Review papers, editorials, letters, comments, and book re-
views were excluded. Reasons for exclusion of reviewed publica-
tions were recorded (Roberts et al., 2006).
2.3. Publication categorization
For included publications, we collected data on afliation of
rst author (categories: academic, government, non-governmental
organization, and private industry), continent in which research
was conducted, and type of taxa studied (mammal, bird, sh,
herpetile, arthropod, and other). We also assessed primary
research topics of the publication, including: (1) animal behavior:
behavior of individual animals, categorized by topics such as
dispersal, home range, habitat selection, mating, nesting, and
25
habituation; (2) population ecology: focused at the population-le-
vel, including topics such as demography, birth, death, nesting, or
reproduction; (3) community ecology: describing interactions be-
tween two or more species; (4) landscape ecology: focused at the
landscape-level, reected in topics such as GIS, landscape pattern
or heterogeneity, edge effects, or land cover; (5) conservation: fo-
cused on wildlife conservation, reected in keywords such as con-
serving, endangered, imperiled, endemic, or rare; (6) human
dimensions: research using surveys or assessment of human atti-
tudes towards wildlife; (7) human-wildlife conict: topics such
as wildlife attacks on humans, economic damage, or animal-vehi-
cle collisions; (8) wildlife management: active management of
wildlife, including strategies such as population control, relocation,
sharpshooting, and sterilization; and (9) wildlife disease: including
both zoonotic and non-zoonotic pathogens harbored by wildlife. In
all instances, multiple options could be selected within a given cat-
egory. Finally, we classied the type of research site where the
study was conducted (i.e., degree of residential development)
based on terms used in the abstract, title, and/or keywords (urban,
suburban, exurban, natural and unknown). While exurban areas
are rural, residential areas beyond the urban fringe (Theobald,
2005), we included them in this study due to their association with
urban habitat. Note that studies conducted only on natural areas
were excluded, but some studies contained both urbanized as well
as natural study sites for comparison. We rened our data collec-
tion process using subsets of 20 publications on which observer
categorizations were compared. Each topic was rened until all ob-
server responses were within 90% agreement (Lawler et al., 2006).
2.4. Non-urban publications
In addition, we randomly selected 150 additional publications
that researched wildlife in non-urban areas and recorded author
afliation, geographic location, focal taxa, and primary research
topics for comparison to our included urban wildlife publications.
To select these studies, 30 of our included urban wildlife articles
were randomly selected within each journal type (animal behavior,
biodiversity conservation, landscape ecology, and wildlife biology),
omitting general science journals as they had insufcient included
urban wildlife articles (n = 2) to compare. For each selected urban
wildlife paper, another literature search was performed on Web
of Science within the selected publications journal and year pub-
lished. Within the search results, we selected the rst research pa-
per that studied wildlife in non-urban areas. If more than one
included urban wildlife publication was selected from the same
journal and year, additional non-urban wildlife research papers
were selected for that journal year.
2.5. Analysis
We summarized the number of urban wildlife publications in-
cluded by our process for each journal and eld (animal behavior,
conservation, ecology, general science, landscape ecology, wildlife
management). We also summarized each as a proportion of the
number of publications by that journal, both overall and by decade.
Similarly, we summarized the proportion of included urban wild-
life papers with various author afliations, continents on which re-
search was conducted, focal taxa, scientic topics of the studies,
and type of study site. We also summarized non-urban papers used
for comparison in the same manner. We used chi-squared tests to
evaluate differences between groups, including testing changes in:
(1) publication rate, author afliation, taxonomic focus, scientic
topic, and research site type between decades; (2) author aflia-
tion, geographic location, taxonomic focus, and scientic topic be-
tween urban and non-urban publications; and (3) taxonomic focus
and geographic location across scientic topics.
26 S.B. Magle et al. / Biological Conservation 155 (2012) 2332

Fig. 1. Percentage of urban wildlife publications from 1971 to 2010 in 16 animal behavior, conservation, ecology, general science, landscape ecology, and wildlife biology
journals. Data are presented as the percentage of the total number of publications published each year by the target journals. Trendline is a second order polynomial function,
indicating increasing publication volume.

3. Results representing 58 studies; Table 1). This was followed by Landscape

We reviewed 1099 total publications generated by our urban
keyword search within the 16 target journals from 1971 to 2010,
ultimately including 571. Of the 528 excluded papers, 300
(56.8%) did not research wildlife, 165 (31.2%) were not research pa-
pers, and 95 (18.0%) did not occur in urban, suburban, or exurban
areas. Note that some papers were excluded on multiple grounds.
3.1. Annual trends
Rates of publication of urban wildlife literature are increasing
(Fig. 1). Before 1991, only 14 papers were included as a result of
our process (1 in 19711980 and 13 in 19811990), representing
0.01% of the publications by our target journals in that period. In
19912000, 128 papers studied wildlife in urban areas, represent-
ing 0.17% of overall publications. By 20012010, the rate of urban
wildlife publications in selected journals had increased to 0.55%,
representing 429 total studies (Fig. 1).
3.2. Journals
Of our target journals, Wildlife Research published the highest
proportion of publications addressing urban wildlife topics (4.1%,
and Urban Planning (3.7%, 85 publications) and Landscape Ecology
(3.2%, 42 publications). However, Biological Conservation contrib-
uted the largest volume of publications (117). The wildlife biology
journals published the highest percentage of urban wildlife publi-
cations during the 1990s but were surpassed in the early 2000s by
the landscape ecology journals (Fig. 2). Publication rates signi-
cantly differed between decades only for the landscape journals
(v2 = 8.46, p = 0.04, df = 3) and not other elds (all v2 < 5.16,
p > 0.18, df = 3). Qualitative trends, however, suggest slow, steady
increases in publication rates in conservation and wildlife journals,
but consistently low rates in animal behavior and ecology journals
(Fig. 2). General science journals were excluded from the analyses
of temporal trends due to exceedingly low number of urban wild-
life papers (n = 2; Table 1, Fig. 2).
3.3. Author afliation
Overall, 73.7% of included urban wildlife publications listed an
academic afliation for their rst author (Table 2). In comparison,
14.5% of lead authors were employed by a state, local, or national
government, 10.9% were afliated with non-governmental organi-
zations, and only 0.70% worked for private companies. Rates did
not signicantly vary between the 1990s and 2000s (all v2 < 1.41,

Table 1
Urban wildlife publications in 16 target journals from 1971 to 2010, as well as total number of publications published over that time period and the percentage that researched
urban wildlife.

Journal type Journal Urban wildlife publications All published publications Percentage
Animal Behavior Animal Behaviour 10 9479 0.11
Behavioral Ecology 11 2211 0.5
Behavioral Ecology and Sociobiology 7 3926 0.18
Conservation American Naturalist 5 5456 0.09
Biological Conservation 117 5237 2.23
Conservation Biology 56 3863 1.45
Ecology Ecology 18 9401 0.19
Ecology Letters 1 1524 0.07
Journal of Applied Ecology 16 3726 0.43
General Science Nature 1 121290 <0.01
Science 1 90350 <0.01
Landscape Ecology Landscape and Urban Planning 85 2305 3.69
Landscape Ecology 42 1301 3.23
Wildlife Biology Journal of Wildlife Management 77 6231 1.24
Wildlife Research 58 1405 4.13
Wildlife Society Bulletin 66 2931 2.25
 S.B. Magle et al. / Biological Conservation 155 (2012) 2332 27

Fig. 2. Percentage of urban wildlife publications from 1971 to 2010 within a given eld, represented by leading high-impact journals.

Table 2
Afliations of rst authors for urban wildlife publications from 1971 to 2010, grouped by decades. Data expressed as percentages of the urban wildlife publications included in our
analyses.

Decade Total % Academic % Government % NGO % Private industry

19711980 1 100.00 0 0 0
19811990 13 69.23 23.08 0 7.69
19912000 128 66.41 20.31 11.72 1.56
20012010 429 75.75 13.28 11.18 0.23
Total 571 73.73 14.52 10.92 0.70

p > 0.16, df = 1), though qualitative trends suggest a slight increase
in work conducted by academics and a decrease in that conducted
by governmental researchers (Table 2). Insufcient publications
were included in the 1980s (13) or 1970s (1) to warrant statistical
analysis.
3.4. Geographic focus
Most urban wildlife studies (291/571, 51.0%) were conducted in
North America, followed by Europe (118/571, 20.7%) and Australia
(105/571, 18.4%). Only 6.8% (39/571) of studies occurred in Asia,
3.7% (21/571) in South America, and 2.8% (16/571) in Africa. Re-
search conducted on more than one continent was rare (9/571,
1.6%).
3.5. Taxonomic focus
The taxa most frequently studied in urban wildlife publications
were birds and mammals, (Table 3), though they were rarely stud-
ied simultaneously (20/571, 3.5%). Arthropods, herpetiles, and sh
were studied less frequently. There were no major shifts in taxa
studied over time; percentages of studies focusing on birds and
mammals were not signicantly different between the 1990s and
2000s (both v2 < 0.19, p > 0.65, df = 1); other taxa and decades
had insufcient data to test for changes. In North America, 40.2%
of urban wildlife publications (117/291) included birds while
40.9% (119/291) studied mammals, as compared to Europe, where
47.5% (56/118) included birds and only 24.6% (29/118) included
mammals, and Australia, with 41.9% bird (44/105) and 52.4% mam-
mal (55/105) studies. The proportion of mammal studies signi-
cantly differed among these continents (v2 = 6.16, p = 0.013,
df = 2), but bird studies did not (v2 = 0.10, p = 0.77, df = 2). Due to
low sample size, we did not test differences among continents
for other focal taxa, include additional continents, or include ear-
lier decades, though a high proportion (53.8%) of the 13 urban
wildlife papers published from 1981 to 1990 focused on mammals
(Table 3).
3.6. Scientic topic
The most frequently represented scientic topics in urban wild-
life research were animal behavior, conservation, and landscape
ecology, followed by wildlife management and population ecology
(Fig. 3). Studies focusing on community ecology (9.3%, 53/571), hu-
man dimensions (8.1%, 46/571), human-wildlife conict (7.2%, 41/
571), disease (2.6%, 15/571), and genetics (2.4%, 14/571) were less
common. The most common topic combination was animal behav-
ior and conservation (42.0%, 240/571), followed by landscape ecol-
ogy and conservation (31.9%, 182/571), animal behavior and
landscape ecology (31.0%, 177/571), animal behavior, landscape
ecology, and conservation (26.1%, 149/571), animal behavior and
wildlife management (18.0%, 103/571), animal behavior and

Table 3
Focal taxa for urban wildlife publications from 1971 to 2011, grouped by decades. Data expressed as percentages of the urban wildlife publications included in our analyses.

Decade Papers % Mammal % Bird % Arthropod % Herpetile % Fish % Other

19711980 1 0 0 0 100.00 0 0
19811990 13 53.85 38.46 0 7.69 7.69 0
19912000 128 36.72 46.09 10.94 6.25 0 4.69
20012010 429 38.20 41.91 11.88 10.22 4.22 4.20
Total 571 38.17 42.73 11.38 9.45 3.32 4.20
28 S.B. Magle et al. / Biological Conservation 155 (2012) 2332
Fig. 3. Percentage of urban wildlife publications from 1971 to 2010 that included various scientic topics. 19912000 and 20012010, which represented >97% of included
publications, are displayed separately.
Fig. 4. Percentage of urban wildlife publications from 1971 to 2010 within scientic topics that included various focal taxa.
population ecology (11.9%, 68/571), and conservation and wildlife
management (10.5%, 60/571). Other combinations were repre-
sented in less than 10% of publications.
Landscape ecology research increased signicantly from the
1990s to the 2000s (Fig. 3, v2 = 6.82, p = 0.009, df = 1) and there
was a trend towards a concurrent decrease in animal behavior re-
search (v2 = 2.89, p = 0.09, df = 1); other topics did not signicantly
differ between decades (all v2 < 0.90, p > 0.34, df = 1). In general,
the proportion of research conducted on different taxa, and on dif-
ferent continents, were similar across topic areas (Figs. 4 and 5),
though a relatively high proportion of urban wildlife management
publications focused on mammals (51.8%, 86/166) and were con-
ducted in North America (62.7%, 104/166). Population ecology
studies of urban wildlife were similarly frequent in North America
(61.3%, 73/119).
3.7. Research site types
The most frequent study site type within our included publica-
tions was urban (83.0%, 474/571). Suburban (15.6%, 89/571) and
exurban studies (4.4%, 25/571) were uncommon. Of the 474 stud-
ies conducted in urban areas, 130 (27.4%) also included research in
natural habitat. Percentages of urban wildlife publications that fo-
cused on urban, suburban, or exurban areas were not signicantly
different between the 1990s and 2000s (all v2 < 1.07, p > 0.30, df =
1).
3.8. Comparing urban and non-urban wildlife publications
The proportion of included studies conducted on urban wildlife
compared to that conducted on wildlife in non-urban systems did
not signicantly differ for any author afliation (all v2 < 0.80,
p > 0.35, df = 1)) or continent (all v2 < 1.32, p > 0.25, df = 1) except
Africa, where there was a slight trend towards a smaller proportion
of urban (2.8%, 16/571) compared to non-urban (8.7%, 13/150)
wildlife studies (v2 = 2.98, p = 0.083, df = 1). Studies on birds also
tended to be more common in urban (42.6%, 243/571) compared
to non-urban (28.7%, 43/150) areas (v2 = 2.75, p = 0.097, df = 1);
the proportion of urban and non-urban wildlife studies did not dif-
fer for any other taxon (all v2 < 2.36, p > 0.12, df = 1). Finally, com-
munity ecology topics were less frequent in urban (9.3%, 53/571)
than in non-urban (29.3%, 44/150) systems (v2 = 10.49, p = 0.001,
df = 1), whereas such proportions did not signicantly differ for
other scientic topics (all v2 < 2.63, p > 0.10).
4. Discussion
The rate of publication of scientic studies of urban wildlife in-
creased more than tenfold in our target journals since 1971, from
less than 0.1% of published articles previous to 1990, to nearly
2% between 2001 and 2010. While only 14 urban wildlife publica-
tions were published before 1991 in the journals we reviewed, we
identied over 400 such studies published between 2000 and
 S.B. Magle et al. / Biological Conservation 155 (2012) 2332
Fig. 5. Percentage of urban wildlife publications from 1971 to 2010 within scientic topics that were conducted on various continents.
2010. Without doubt, urban wildlife research is a growing eld,
both in terms of number of articles published and in the proportion
of published work that focuses on urban areas. Similarly, in a pre-
vious review of two wildlife biology journals (Journal of Wildlife
Management and Wildlife Society Bulletin), Adams (2005) found that
the proportion of studies on urban wildlife increased from 12% in
the early 1980s to 34% in the late 1990s and early 2000s (Adams,
2005). This developing focus likely reects the identication of an
important gap in ecological research (Adams et al., 1987; Chace
and Walsh, 2006; Marzluff et al., 2008) as well as mounting recog-
nition of the rapid expansion of urban areas coupled with an over-
all decrease of available alternate habitat for wildlife (Dearborn
and Kark, 2010; McKinney, 2006). Indeed, we note the emergence
of a suite of urban ecology journals that publish wildlife studies
(e.g., Cities and the Environment, Landscape and Urban Planning, Ur-
ban Ecology, Urban Ecosystems), as well as journals that handle hu-
man-wildlife conict papers specically, including those in urban
areas (e.g., Human-Wildlife Conicts, Human Dimensions in Wildlife).
These specic journals failed to meet our selection criteria of high-
impact journals published before 1990 and thus were not included
in our analyses, but nonetheless represent important additional
outlets for urban wildlife research.
While the trend is increasing, we emphasize that overall rates of
publication of urban wildlife research remain low (<2% in our tar-
get journals), particularly considering that urban sprawl and its
impacts on biodiversity are accelerating globally. Indeed, urbaniza-
tion is a dominant demographic trend (Marzluff et al., 2008) and
the planet is undergoing an unprecedented urban transition
(Ramalho and Hobbs, 2012); for the rst time in human history,
the majority of humans on the planet live in urban areas, with
nearly all population growth this century expected to be urban
(United Nations Population Fund, 2007; United Nations, 2010).
Urbanization has become a primary agent of habitat loss and frag-
mentation, and as a result, is also a leading cause of species endan-
germent globally (Czech et al., 2000; Mcdonald et al., 2008).
Biologists have not emphasized urban study systems for a variety
of reasons, including a traditional preference to focus on native
species in relatively remote, pristine, and undisturbed ecosystems,
as well as logistical, political, and nancial obstacles to urban re-
search (Miller and Hobbs, 2002).
Not only were urban wildlife studies relatively infrequent, but
they also varied considerably among disciplines. The landscape
ecology journals published urban wildlife research in the highest
proportion, though it should be noted we included only two such
29
journals, and one of those, Landscape and Urban Planning, clearly fo-
cuses on urban areas. Wildlife biology and then conservation jour-
nals published the next highest percentage of their articles on
urban wildlife, with the greatest absolute volume of such studies
published in Biological Conservation. Miller and Hobbs (2002) noted
that fewer than 6% of the publications published in the journal
Conservation Biology between 1995 and 1999 included work in ur-
ban, suburban, or exurban areas; our own more exhaustive review
in that journal suggests lower rates (<2% overall) for wildlife stud-
ies specically. In a similar review, Collins et al. (2000) found that
only 0.4% of papers published in 9 leading ecological journals be-
tween 1995 and 2000 focused on urban species (Collins et al.,
2000). Excluding landscape ecology, we also recorded very low
rates of publication of urban wildlife research in our three target
ecological journals (0.2% overall), providing no evidence for appar-
ent increase since 2000. Our literature review demonstrated that
such studies were rare in animal behavior journals as well, and
were exceedingly uncommon in the high-impact general science
journals. Virtually all reviews on this topic conclude that an in-
creased focus on urban systems is needed among ecologists (e.g.,
Adams, 2005; Chace and Walsh, 2006; Collins et al., 2000; Gehrt
et al., 2010; Mcdonald et al., 2008; Marzluff et al., 2008; Miller
and Hobbs, 2002; Pickett et al., 2008). Although our results do indi-
cate some progress in this direction, such publications are uneven
among disciplines and remain relatively scarce in our target
journals.
The majority (ca. 75%) of published urban wildlife research has
been directed by academics, as assessed by rst-authorship of
resulting publications. While studies led by government research-
ers represent the next most common type of urban wildlife study,
it is far less common (ca. 15%). Our comparisons across decades
suggests that this disparity, which was noted in the 1980s (Adams
et al., 1987) and also is evident between the wildlife management
and conservation elds (Jensen and Krausman, 1993), is not dimin-
ishing and may even be widening. Our results also suggest that this
disparity is similar for both urban and non-urban wildlife studies.
As publications represent the primary currency for advancement in
academic careers, it is logical to expect academics to publish more
frequently than other researchers in this eld. However, linking ba-
sic knowledge to informed practice has been identied as one of
the greatest challenges to the discipline of urban ecology (Marzluff
et al., 2008). Moreover, one of the stated goals of urban wildlife re-
search is to include non-academic researchers in the production of
knowledge (Young and Wolf, 2006), and governmental and non-
30 S.B. Magle et al. / Biological Conservation 155 (2012) 2332
governmental agencies may bring different perspectives to such
research and hence may provide novel or unique insights. Our
study design focused on primary authors, and as such it is possible
that non-academic researchers are becoming more involved in ur-
ban wildlife research, either as non-primary authors or at a level
typically recognized in acknowledgments. Young and Wolf
(2006) identied a trend towards more non-academic research in
the purely urban journals Urban Ecology and Urban Ecosystems be-
tween the periods 19751986 and 19972004, which is encourag-
ing, but our results reveal little evidence for such a trend for urban
wildlife publications in the primary animal behavior, conservation,
and ecology literature.
In terms of geographic focus, more than half (51.0%) of the ur-
ban wildlife studies we reviewed were conducted in North Amer-
ica, a proportion similar to wildlife studies conducted in non-
urban areas. Certainly it should be noted that all of our selected
journals were English-language journals, and most are themselves
based in North America, and this may impact our results. However,
all included journals accept international work, and are among the
leading journals in their eld, so we believe this result reects a
true focus in current research. The relatively large number of stud-
ies in Europe (20.6%) is expected given that Europe was the rst
continent to begin urban ecological surveys (Adams, 2005). The
substantial representation of Australia (18.5% of studies) is likely
partially a result of Australias recent focus on urban wildlife ecol-
ogy (Adams, 2005), but perhaps also was inuenced by our inclu-
sion of Wildlife Research, an international journal based in
Australia. When that journal is omitted, however, the proportion
of urban wildlife studies conducted in Australia drops to 9.3%,
which still exceeds all other continents but North America and Eur-
ope. The small number of studies published from Asia, South Amer-
ica, and Africa are particularly worrisome given that these areas
are experiencing rapid human population growth and resulting
urbanization (Chace and Walsh, 2006) and contain numerous bio-
diversity hotspots (Myers et al., 2000; Mcdonald et al., 2008); in
Africa specically, wildlife studies tended to be less common in ur-
ban compared to non-urban systems. For some studies, a language
barrier may have prevented researchers in these areas from pub-
lishing in the top journals, and funding restrictions may have lim-
ited the scope of such studies to the point where publication in
high-impact journals was challenging. Finally, comparative studies
on more than one continent are rare. While logistically difcult,
such research will likely be necessary for discovery and renement
of global urban ecological principles.
Birds were the most frequently studied taxon, consistent with
prior reviews (Adams, 2005; Marzluff et al., 2001), and were also
more frequently studied in urban compared to non-urban areas,
though mammals were also frequent targets for urban ecological
research, particularly in North America and Australia. Taxonomic
focus did not appear to differ between the 1990s and 2000s,
although of the few urban wildlife papers before 1990, the majority
focused on mammals. Herpetiles, sh, and arthropods were infre-
quent focal taxa of urban wildlife studies and thus represent taxo-
nomic gaps that warrant increased attention in future research
efforts (see also McIntyre, 2000; Paul and Meyer, 2001; Shochat
et al., 2004). However, it should be noted that some journals used
in this analysis (i.e. Journal of Wildlife Management, Wildlife Society
Bulletin) deliberately exclude manuscripts with invertebrates and
sh as main subjects, thus limiting the appearance of these taxa.
Amphibians (Hamer and Parris, 2011) and aquatic invertebrates
(Paul and Meyer, 2001) are particularly susceptible to urban im-
pacts, including pollutants originating in urban areas, and may rep-
resent excellent focal species for studies of ecotoxicology or urban
adaptation. Arthropods also represent the majority of wildlife spe-
cies on Earth (Wilson, 1992) and are critical for ecosystem function
(Bolger et al., 2000), including providing important services for hu-
mans such as pollination and biological control (Losey and Vaugh-
an, 2006); additional study is necessary to fully understand how
these functions are altered in urban landscapes and the implica-
tions of these alterations. Less is known about sh responses to
urbanization compared to other taxa, but available studies suggest
that native sh abundance and diversity tend to decline, and inva-
sive species increase, in more urbanized areas (Paul and Meyer,
2001; Wolter, 2008). Finally, studies investigating multiple taxa,
and even more than simply one species, were uncommon, as evi-
denced by the low proportion (9.3%) of studies investigating com-
munity ecology. Wildlife studies incorporating multiple species
and community dynamics were more common outside of urban
systems, and these could help guide similar studies in metropoli-
tan areas. Assessing interactions between organisms will be neces-
sary to better understand ecological dynamics of urban systems
(e.g. Crooks and Soul, 1999; Faeth et al., 2005; Crooks et al., 2010).
Urban ecology is an inherently interdisciplinary eld (Marzluff
et al., 2008), and this was reected in the diversity of research top-
ics addressed in urban wildlife studies. The most common topic for
publications in our target journals was animal behavior, with the
majority of these studies focusing on habitat requirements, move-
ment, or dispersal within urban settings. Many studies have inves-
tigated behavioral adaptations of wildlife to urban areas, including
changes to activity levels, foraging behavior, and reproduction
(Ditchkoff et al., 2006). However, despite the prevalence of studies
assessing the behavior of urban wildlife, the animal behavior jour-
nals generally published a smaller proportion of urban studies than
the conservation or wildlife journals, suggesting that the wider
community of animal behavior scientists, many motivated by the-
oretical and not applied questions (Angeloni et al., 2008), are not
yet fully engaged in urban research. Studies regarding the conser-
vation of wildlife in urban areas were next most common. Conserv-
ing species in areas of high human density can be controversial, as
some of these urban species may be invasive, harbor disease, and
conict with humans (Angermeier, 1994; Forman et al., 2003; Ur-
banek et al., 2011). Nevertheless, conservation of urban wildlife
has numerous benets, including protecting rare and threatened
species, understanding wildlife response to environmental change
such as global warming, and providing ecosystem services and
education to humans (Dearborn and Kark, 2010). The relatively
new discipline of landscape ecology also has been increasingly ap-
plied to urban wildlife (see also Adams, 2005), partly because the
large spatial extent of urban areas are directly applicable to
broad-scale landscape ecology approaches. Moreover, landscape
ecology also explicitly recognizes landscape context and inter-con-
nectedness of systems (Wu, 2008), which can be powerfully ap-
plied to urban areas that are characterized by complex ows
between ecological, socio-economic, and physical processes. Stud-
ies on the population ecology of urban species previously have
been identied as sparse in wildlife biology journals (Adams,
2005), but a relatively large proportion of our studies (20.9%), as-
sessed in a broader range of journals, investigated this topic. Less
represented were studies of genetics, disease, or human-wildlife
conict such as animal-vehicle collisions, landscape damage, or
pet predation, though this has been cited as a growing eld of re-
search (Adams, 2005). It is notable that while direct assessments of
wildlife harm to humans and property were rare, studies of man-
agement of wildlife, sometimes with the stated goal of reducing
these conicts, were much more common. This disconnect may
suggest that the degree of damage caused by wildlife should be di-
rectly assessed more often to determine if management action is
warranted.
In our review, we included studies conducted across a range of
residential land use types, including urban, suburban, and exurban
systems. Urban systems, characterized by high density develop-
ment, dominated the urban wildlife literature (ca. 83% of studies),
 S.B. Magle et al. / Biological Conservation 155 (2012) 2332
as would be expected given our focus on urban studies. Suburban
areas, land use dominated by lower density residential develop-
ment, are widespread and characterized by generalist and invasive
species and growing conict between humans and wildlife
(DeStefano and DeGraff, 2003). Nonetheless, suburban wildlife
studies were relatively uncommon (ca. 15%) and we did not detect
any apparent increase in their frequency over time. Exurban re-
search in residential development beyond the urban fringe was
rare, perhaps partly because such regions have been identied as
important for study only recently (Theobald, 2001). However,
these rural residential areas are critical to ecology and conserva-
tion because they are frequently located adjacent to protected nat-
ural areas and are expanding even more rapidly than urban areas
in many regions (Theobald, 2005). Additional wildlife research in
suburban and exurban areas therefore would be valuable.
5. Conclusions
Urban wildlife biology has frequently been cited as an emergent
discipline (e.g. Adams, 2005; Gehrt et al., 2010; Werner, 2011). Our
review of 16 leading journals from the animal behavior, conserva-
tion, ecology, general science, landscape ecology, and wildlife biol-
ogy elds conrms that the rates at which journals are publishing
urban wildlife studies are increasing. However, the expansion of
urban wildlife literature is varied, with animal behavior, ecology,
and general science journals disciplines lagging behind those in
landscape ecology, wildlife biology, and conservation. Moreover,
the overall rates of publications of such studies remain quite low,
particularly when considering the extent of urban growth and its
impact on biodiversity globally. By analogy, numerous barriers ex-
ist between researchers of animal behavior and conservation, with
animal behavior scientists potentially viewing the more applied
subject of conservation as less objective, less rigorous, or insuf-
ciently challenging (Angeloni et al., 2008). It is conceivable that
similar objections deter basic scientists from engaging in urban
wildlife research, particularly given that studies in such human-
dominated landscapes frequently generate applied questions as
opposed to the fundamental scientic research more characteristic
of natural or pristine systems. However, animal behavioral princi-
ples were frequently applied to urban studies we reviewed, even
though those studies were rarely published in behavioral journals.
These discrepancies would seem to represent an opportunity for
behavioral and ecological researchers, and their target journals,
to seize upon the growing phenomena of urbanization for future
study. Similarly, non-academic researchers were infrequent rst
authors of urban wildlife work, and thus represent an underex-
ploited yet valuable resource in this eld.
A disproportionate amount of urban wildlife research is pub-
lished from North America, Europe, and Australia, even though
South America, Africa, and Asia are rapidly urbanizing and support
much of the earths biodiversity (Mcdonald et al., 2008); this rep-
resents an important gap in our knowledge that should be ad-
dressed. Similarly, most urban research is conducted on birds
and mammals, with herpetiles, sh, and arthropods relatively
unrepresented. Although charismatic vertebrates play important
roles both ecologically and economically, studies of less under-
stood taxa in urban areas should be encouraged. We also expected
an increase in studies conducted in suburban and exurban areas
due to the rapid expansion of such development and research indi-
cating its importance (DeStefano and DeGraff, 2003; Theobald,
2001, 2005), yet we could not detect any such increase within
our target journals. This gap represents another potentially impor-
tant avenue for wildlife research in the future.
Overall, knowledge regarding urban wildlife and their adapta-
tions to, and persistence within, human-dominated landscapes will
31
continue to advance. At present, however, such advancement re-
mains uneven, with important research gaps to be resolved. Urban
wildlife research is still a young eld with unfullled potential and
numerous opportunities for a wide range of scientists from a diver-
sity of disciplines. We contend that the interdisciplinary science of
understanding the animal species who share our cities has tremen-
dous and increasing importance for our urbanizing planet.
Acknowledgments
We thank L. Lehrer for assistance with reviews of publications.
We also thank S. Gehrt and an anonymous reviewer for helpful
comments on the manuscript. Funding for VMH, SBM, and MV
was provided by the Lincoln Park Zoo and the Davee Foundation.
References
Adams, L.W., Leedy, D.L., McComb, W.C., 1987. Urban wildlife research and
education in North American colleges and universities. Wildlife Society
Bulletin 15, 591595.
Adams, L.W., 2005. Urban wildlife ecology and conservation: a brief history of the
discipline. Urban Ecosystems 8, 139156.
Angeloni, L., Schlaepfer, M.A., Lawler, J.J., Crooks, K.R., 2008. A reassessment of the
interface between conservation and behavior. Animal Behaviour 75, 731737.
Angermeier, P.L., 1994. Does biodiversity include articial diversity? Conservation
Biology 8, 600602.
Australian Research Centre for Urban Ecology, 2011. Australian Research Centre for
Urban Ecology. <http://arcue.botany.unimelb.edu.au/>.
Blair, R.B., Johnson, E.M., 2008. Suburban habitats and their role for birds in the
urbanrural habitat network: points of local invasion and extinction?
Landscape Ecology 23, 11571169.
Bolger, D.T., Alberts, A.C., Sauvajot, R.M., Potenza, P., McCalvin, C., Tran, D., Mazzoni,
S., Soul, M.E., 1997. Response of rodents to habitat fragmentation in coastal
southern California. Ecological Applications 7, 552563.
Bolger, D.T., Suarez, A.V., Crooks, K.R., Morrison, S.A., Case, T.J., 2000. Arthropods in
urban habitat fragments in southern California: area, age, and edge effects.
Ecological Applications 10, 12301248.
Bolitzer, B., Netusil, N.R., 2000. The impact of open spaces on property values in
Portland, Oregon. Journal of Environmental Management 59, 185193.
Bradley, C.A., Altizer, S., 2007. Urbanization and the ecology of wildlife diseases.
Trends in Ecology and Evolution 22, 95102.
Chace, J.F., Walsh, J.J., 2006. Urban effects on native avifauna: a review. Landscape
and Urban Planning 74, 4669.
Chawla, L., 1999. Life paths into effective environmental action. Journal of
Environmental Education 31, 1526.
Collins, J.P., Kinzig, A., Grimm, N.B., Fagan, W.F., Hope, D., Wu, J., Borer, E.T., 2000. A
new urban ecology: modeling human communities as integral parts of
ecosystems poses special problems for the development and testing of
ecological theory. American Scientist 88, 416425.
Crooks, K.R., Soul, M.E., 1999. Mesopredator release and avifaunal extinctions in a
fragmented system. Nature 400, 563566.
Crooks, K.R., Suarez, A.V., Bolger, D.T., 2004. Avian assemblages along a gradient of
urbanization in a highly fragmented landscape. Biological Conservation 115,
451462.
Crooks, K.R., Riley, S.P.D., Gehrt, S.D., Gosselink, T.E., Van Deelen, T.R., 2010.
Community ecology of urban carnivores. In: Gehrt, S.D., Riley, S.P.D., Cypher,
B.L. (Eds.), Urban Carnivores: Ecology, Conict, and Conservation. John Hopkins
University Press, Baltimore, MD, USA, pp. 185196.
Curtis, P.D., Hadidian, J., 2010. Responding to humancarnivore conicts in urban
areas. In: Urban Carnivores: Ecology, Conict, and Conservation. Johns Hopkins
University Press, Baltimore, MD, USA, pp. 201212.
Czech, B., Krausman, P.R., Devers, P.K., 2000. Economic associations among causes of
species endangerment in the United States. BioScience 50, 593601.
Dearborn, D.C., Kark, S., 2010. Motivations for conserving urban biodiversity.
Conservation Biology 24, 432440.
DeStefano, S., DeGraff, R.M., 2003. Exploring the ecology of suburban wildlife.
Frontiers in Ecology and the Environment 1, 95101.
Ditchkoff, S.S., Saalfeld, S.T., Gibson, C.J., 2006. Animal behavior in urban
ecosystems: modications due to human-induced stress. Urban ecosystems 9,
512.
Faeth, S.H., Warren, P.S., Shochat, E., Marussich, W.A., 2005. Tropic dynamics in
urban communities. BioScience 55, 399407.
FitzGibbon, S.I., Putland, D.A., Goldizen, A.W., 2007. The importance of functional
connectivity in the conservation of a ground-dwelling mammal in an urban
Australian landscape. Landscape Ecology 22, 15131525.
Forman, R.T.T., Sperling, D., Bissonette, J.A., Clevenger, A.P., Cutshall, C.D., Dale, V.H.,
Fahrig, L., France, R., Goldman, C.R., Heanue, K., Jones, J.A., Swanson, F.J.,
Turrentine, T., Winter, T.C., 2003. Road Ecology: Science and Solutions. Island
Press, Washington, DC.
Forman, R.T.T., 2008. The urban region: natural systems in our place, our
nourishment, our home range, our future. Landscape Ecology 23, 251253.
32 S.B. Magle et al. / Biological Conservation 155 (2012) 2332
Frankham, R., 2006. Genetics and landscape connectivity. In: Crooks, K.R., Sanjayan,
M. (Eds.), Connectivity Conservation. Cambridge University Press, United
Kingdom, pp. 7296.
Gehrt, S.D., 2010. The urban ecosystem. In: Urban Carnivores: Ecology, Conict, and
Conservation. Johns Hopkins University Press, Baltimore, MD, USA, pp. 311.
Gehrt, S.D., Riley, S.P.D., 2010. Coyotes (Canis latrans). In: Urban Carnivores:
Ecology, Conict, and Conservation. Johns Hopkins University Press, Baltimore,
MD, USA, pp. 7998.
Gehrt, S.D., Riley, S.P.D., Cypher, B.L. (Eds.), 2010. Urban Carnivores: Ecology,
Conict, and Conservation. Johns Hopkins University Press, Baltimore, MD, USA.
Gibb, H., Hochuli, D.F., 2002. Habitat fragmentation in an urban environment: large
and small fragments support different arthropod assemblages. Biological
Conservation 106, 91100.
Grimm, N.B., Grove, J.M., Pickett, S.T.A., Redman, C.L., 2000. Integrated approaches
to long-term studies of urban ecological systems. BioScience 50, 571584.
Hamer, A.J., Parris, K.M., 2011. Local and landscape determinants of amphibian
communities in urban ponds. Ecological Applications 21, 378390.
Jensen, M.N., Krausman, P.R., 1993. Conservation biologys literature: new wine or
just a new bottle? Wildlife Society Bulletin 21, 199203.
Lawler, J.J., Aukeme, J.E., Grant, J.B., Halpern, B.S., Kareiva, P., Nelson, C.R., Ohleth, K.,
Olden, J.D., Schlaepfer, M.A., Silliman, B.R., Zaradic, P., 2006. Conservation
science. A 20-year report card. Frontiers in Ecology and the Environment 4,
473480.
Losey, J.E., Vaughan, M., 2006. The economic value of ecological services provided
by insects. BioScience 56, 311323.
Magle, S.B., Theobald, D.T., Crooks, K.R., 2009. Comparing isolation metrics
predicting the distribution of a highly interactive species across an urban
gradient. Landscape Ecology 24, 267280.
Marzluff, J.M., Bowman, R., Donnelly, R. (Eds.), 2001. Avian Ecology and
Conservation in an Urbanizing World. Kluwer Academic Publishers, Boston,
Massachusetts, USA.
Marzluff, J.M., Schulenberger, E., Endlicher, W., Alberti, M., Bradley, G., Ryan, C.,
Simon, U., ZumBrunnen, C. (Eds.), 2008. Urban Ecology: An International
Perspective on the Interaction Between Humans and Nature. Springer, New
York, New York, USA.
Mayer, P., 2010. Urban ecosystems research joins mainstream ecology. Nature 467,
153.
Mcdonald, R.I., Kareiva, P., Forman, R.T.T., 2008. The implications of current and
future urbanization for global protected areas and biodiversity conservation.
Biological Conservation 141, 16951703.
McIntyre, N.E., 2000. Ecology of urban arthropods: a review and a call to action.
Annals of the Entomological Society of America 93, 825.
McKinney, M.L., 2002. Urbanization, biodiversity, and conservation. Bioscience 52,
883890.
McKinney, M.L., 2006. Urbanization as a major cause of biotic homogenization.
Biological Conservation 127, 247260.
Mendes, W., Balmer, K., Kaethler, T., Rhoads, A., 2008. Using land inventories to plan
for urban agriculture: experiences from Portland and Vancouver. Journal of the
American Planning Association 74, 435449.
Miller, J.R., Hobbs, R.J., 2002. Conservation where people live and work.
Conservation Biology 16, 330337.
Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B., Kent, J., 2000.
Biodiversity hotspots for conservation priorities. Nature 403, 853858.
Nowak, D.J., Crane, D.E., Stevens, J.C., 2006. Air pollution removal by urban trees and
shrubs in the United States. Urban Forestry and Urban Greening 4, 115123.
View publication stats
Orr, W., 1993. Love it or lost it: the coming biophilia revolution. In: Kellert, S.R.,
Wilson, E.O. (Eds.), The Biophilia Hypothesis. Island Press, Washington, DC, pp.
415440.
Paul, M.J., Meyer, J.L., 2001. Streams in the urban landscape. Annual Review of
Ecology and Systematics 32, 333365.
Pickett, S.T.A., Cadenasso, M.L., Grove, J.M., Groffman, P.M., Band, L.E., Boone, C.G.,
Burch Jr., W.R., Grimond, C.S.B., Hom, J., Jenkins, J.C., Law, N.L., Nilon, C.H.,
Pouyat, R.V., Szlavecz, K., Warren, P.S., Wilson, M.A., 2008. Beyond urban
legends: an emerging framework of urban ecology, as illustrated by the
Baltimore ecosystem study. BioScience 58, 139150.
Ramalho, C.E., Hobbs, R.J., 2012. Time for a change: dynamic urban ecology. Trends
in Ecology and Evolution 27, 179188.
Roberts, P.D., Stewart, G.B., Pullin, A.S., 2006. Are review articles a reliable source of
evidence to support conservation and environmental management? A
comparison with medicine. Biological Conservation 132, 409423.
Shochat, E., Stefanov, W.L., Whitehouse, M.E.A., Faeth, S.H., 2004. Urbanization and
spider diversity: inuences of human modication of habitat structure and
productivity. Ecological Applications 14, 268280.
Simberloff, D., 1997. The biology of invasions. In: Simberloff, D., Schmitz, D.C.,
Brown, T.C. (Eds.), Strangers in Paradise. Island Press, Washington, DC, pp. 319.
Sukopp, H., 2002. On the early history of urban ecology in Europe. Prelia Praha 74,
373393.
Taylor, L.H., Latham, S.M., Woolhouse, M.E., 2001. Risk factors for human disease
emergence. Philosophical Transactions of the Royal Society B: Biological
Sciences 356, 983989.
Theobald, D.M., 2001. Land use dynamics beyond the American urban fringe.
Geographical Review 91, 544564.
Theobald, D.M., 2005. Landscape patterns of exurban growth in the USA from 1980
to 2020. Ecology and Society 10, 32 (online).
United Nations, 2010. World Urbanization Prospects. The 2009 Revision, Highlights,
United Nations.
United Nations Population Fund. 2007. State of the World Population 2007.
Unleashing the Potential of Urban Growth. United Nations Population Fund.
Urbanek, R.E., Allen, K.R., Nielsen, C.K., 2011. Urban and suburban deer management
by state wildlife-conservation agencies. Wildlife Society Bulletin 35, 310
315.
Waddell, P., Moore, T., 2008. Forecasting demand for urban land. In: Marzluff, J.M.,
Schulenberger, E., Endlicher, W., Alberti, M., Bradley, G., Ryan, C., ZumBrunnen,
C., Simon, U. (Eds.), Urban Ecology: An International Perspective on the
Interaction Between Humans and Nature. Springer Science, New York, NY,
USA, pp. 493518.
Werner, P., 2011. The ecology of urban areas and their functions for species
diversity. Landscape and Ecological Engineering 7, 231240.
Wilson, E.O., 1992. The Diversity of Life. Penguin Press, London, UK.
Wolter, C., 2008. Towards a mechanistic understanding of urbanizations impacts on
sh. In: Marzluff, J.M., Schulenberger, E., Endlicher, W., Alberti, M., Bradley, G.,
Ryan, C., ZumBrunnen, C., Simon, U. (Eds.), Urban Ecology: An International
Perspective on the Interaction Between Humans and Nature. Springer Science,
New York, NY, USA, pp. 425436.
Wu, J., 2008. Making the case for landscape ecology: an effective approach to urban
sustainability. Landscape Journal 27, 4150.
Young, R.F., Wolf, S.A., 2006. Goal attainment in urban ecology research: a
bibliometric review 19752004. Urban Ecosystems 9, 179193.