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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


and abundance are often quite high (Chace and Walsh, 2006;
The rate at which the world is urbanized is accelerating (Dear- McKinney, 2006). Urban biota increasingly is comprised of non-na-
born and Kark, 2010; Forman, 2008; McKinney, 2002; Pickett et al., tive species that are both directly imported and benetted by ur-
2008; Ramalho and Hobbs, 2012), and perhaps as a result, interest ban land modication (McKinney, 2006; Simberloff, 1997).
in wildlife in urban areas appears to be growing (Adams, 2005; For- Animals that reside in cities frequently conict with humans,
man, 2008; Mayer, 2010; Mcdonald et al., 2008; Miller and Hobbs, including incidents of animal-vehicle collision (Forman et al.,
2002). Despite some study of urban plant species and a few iso- 2003), animal attacks on pets (Curtis and Hadidian, 2010; Gehrt
lated investigations of urban wildlife in Europe, systematic re- and Riley, 2010), and damage to lawns and landscaping (Urbanek
search on animal species in and around cities did not begin in et al., 2011). In addition, urban areas, characterized by high densi-
earnest until the 1970s (Gehrt, 2010; Sukopp, 2002). Since that ties of both humans and wildlife, represent a signicant risk for the
time, urban ecology has been formalized as an international, inter- spread of disease (Bradley and Altizer, 2007). Zoonotic pathogens,
disciplinary eld (e.g., Marzluff et al., 2008), with several journals diseases that can spread from wildlife to humans, now constitute
devoted solely to the topic, and research boosted by several 75% of all emerging human diseases (Taylor et al., 2001).
high-prole government projects such as the urban Long-Term To date, no review has provided a quantitative and comprehen-
Ecological Research sites in the United States (Grimm et al., sive evaluation of long-term trends in urban wildlife research
2000), the urban group of the United Nations Educational, Scien- across a diversity of scientic disciplines and their associated jour-
tic, and Cultural Organizations Man and the Biosphere program nals. Herein, we adopt a systematic review process to evaluate
(Adams, 2005), and the Australian Research Centre for Urban Ecol- trends in urban wildlife research over time and in different scien-
ogy (ARCUE, Australian Research Centre for Urban Ecology, 2011). tic elds, and also assess geographic (Chace and Walsh, 2006) and
Despite this emergent eld, however, relatively little is known taxonomic (Lawler et al., 2006) focus, including identifying gaps
about historic and recent trends in urban wildlife research. where urban wildlife research is scarce. Given recent attention
Urban ecosystems are not only increasingly common, but they on urbanization and its impacts, we expected a rapidly increasing
can play important ecological roles (Dearborn and Kark, 2010). This number of scientic publications focused on urban wildlife. We
may include protection of local biodiversity and rare or imperiled also expected that birds would represent the majority of urban
species (Bolger et al., 1997; Crooks et al., 2004), either directly wildlife studies (following Adams, 2005; Marzluff et al., 2001),
for organisms that reside in urban areas or via creation of corridors and that such research would be biased towards North America
or stepping stone patches for individuals moving through cities and Europe given extensive urban growth and an associated focus
(Gibb and Hochuli, 2002; FitzGibbon et al., 2007). Moreover, in on urban research. We also investigated author afliation, predict-
addition to their role in ecology and conservation, urban green ing that while academics conduct the majority of urban wildlife
space provides important benets to humans. These benets in- research, studies directed by governmental agencies and non-gov-
clude ecosystem services such as pollination (Mendes et al., ernmental organizations may be increasing as this is a stated goal
2008), air quality improvement (Nowak et al., 2006), and carbon in urban ecology research (Young and Wolf, 2006). Further, we
sequestration (Pickett et al., 2008), but also extend to simple evaluated studies conducted across a gradient of residential land
enhancement of property value (Bolitzer and Netusil, 2000; use types, including urban, suburban, and exurban systems,
Waddell and Moore, 2008). Perhaps one of the most powerful roles expecting an increase in suburban and exurban studies given
of urban ecosystems is their ability to connect urban citizens with heightened interest in development impacts outside of high-den-
their environment. It is often difcult for residents to develop an sity urban areas (DeStefano and DeGraff, 2003; Theobald, 2001,
appreciation for wildlife when conservation messages and pro- 2005). Finally, we examined major research topics in urban wildlife
grams focus on distant species or places (Orr, 1993). Urban habitats studies to identify primary scientic questions addressed by such
allow people to engage directly with nature where they live and projects, and also compared such studies to those conducted in
recreate, and this approach may be particularly effective for nature non-urban systems. Our study reveals potential gaps in current
education and awareness (Chawla, 1999; Dearborn and Kark, work, illustrates areas of strength and weakness, and provides
2010). The rate at which natural habitats are conserved for wildlife guidance for future urban wildlife research efforts.
is showing signs of decline (Blair and Johnson, 2008), and conse-
quently urban conservation areas will assume greater importance
in the future. 2. Materials and methods
Urban areas are also characterized by their own unique chal-
lenges. Fragmented urban landscapes isolate natural populations Our methods followed recommendations of Roberts et al.
and restrict dispersal and functional connectivity, which can lead (2006) for systematic reviews, although as we were assessing
to demographic and genetic instabilities (Frankham, 2006; Magle trends as opposed to the effect of a process, policy, or intervention,
et al., 2009) and community-level impacts (Crooks and Soul, some components of the systematic review process were not appli-
1999; Faeth et al., 2005; Crooks et al., 2010). Biotic communities cable. Specically, as we were interested in study focus rather than
S.B. Magle et al. / Biological Conservation 155 (2012) 2332 25

outcomes, determining a hierarchy of evidence by which to assign habituation; (2) population ecology: focused at the population-le-
weight to publications was unnecessary. vel, including topics such as demography, birth, death, nesting, or
reproduction; (3) community ecology: describing interactions be-
2.1. Journal selection tween two or more species; (4) landscape ecology: focused at the
landscape-level, reected in topics such as GIS, landscape pattern
First, we identied up to three target journals within the animal or heterogeneity, edge effects, or land cover; (5) conservation: fo-
behavior, conservation, ecology, landscape ecology, and wildlife cused on wildlife conservation, reected in keywords such as con-
biology elds. We used Journal Citation Reports within the Insti- serving, endangered, imperiled, endemic, or rare; (6) human
tute for Scientic Information (ISI) Web of Science (Thomson dimensions: research using surveys or assessment of human atti-
Reuters, New York, NY, USA) to rank journals based on impact tudes towards wildlife; (7) human-wildlife conict: topics such
factor in 2000, using ISIs standardized subject categories of behav- as wildlife attacks on humans, economic damage, or animal-vehi-
ioral science (for behavior journals), biodiversity conservation (for cle collisions; (8) wildlife management: active management of
conservation journals), and ecology (for ecology, landscape ecol- wildlife, including strategies such as population control, relocation,
ogy, and wildlife biology journals). We chose the year 2000 as it sharpshooting, and sterilization; and (9) wildlife disease: including
represented an approximate midpoint in the period where we both zoonotic and non-zoonotic pathogens harbored by wildlife. In
anticipated high rates of urban wildlife publication (1990-present). all instances, multiple options could be selected within a given cat-
Within the ecology topic, we dened a wildlife biology journal as a egory. Finally, we classied the type of research site where the
journal with wildlife in the title, and a landscape ecology journal study was conducted (i.e., degree of residential development)
as a journal with landscape in the title. We omitted journals that based on terms used in the abstract, title, and/or keywords (urban,
did not publish at least 50 articles in 2000 and those that did not suburban, exurban, natural and unknown). While exurban areas
exist prior to 1990 to ensure adequate sample size for analysis, are rural, residential areas beyond the urban fringe (Theobald,
and also omitted any journal that, based on its title, could not rea- 2005), we included them in this study due to their association with
sonably be expected to regularly and frequently publish on wildlife urban habitat. Note that studies conducted only on natural areas
(e.g., Molecular Ecology, Behavioral and Brain Sciences). We selected were excluded, but some studies contained both urbanized as well
up to three journals from those that remained in each eld based as natural study sites for comparison. We rened our data collec-
on the highest impact factor ratings; we selected the highest-tier tion process using subsets of 20 publications on which observer
journals because they have the highest readership, and thus best categorizations were compared. Each topic was rened until all ob-
reect exposure of the scientic community to the topic. Following server responses were within 90% agreement (Lawler et al., 2006).
these criteria, the journals selected from each ISI subject category
were: Animal Behaviour, Behavioral Ecology, and Behavioral Ecology 2.4. Non-urban publications
& Sociobiology for animal behavior; American Naturalist, Biological
Conservation, and Conservation Biology for biodiversity conserva- In addition, we randomly selected 150 additional publications
tion; Ecology, Ecology Letters, and Journal of Applied Ecology for ecol- that researched wildlife in non-urban areas and recorded author
ogy; and Journal of Wildlife Management, Wildlife Research, and afliation, geographic location, focal taxa, and primary research
Wildlife Society Bulletin for wildlife biology. In the landscape ecol- topics for comparison to our included urban wildlife publications.
ogy eld, only two journals met our criteria, Landscape Ecology To select these studies, 30 of our included urban wildlife articles
and Landscape and Urban Planning. For comparative purposes, we were randomly selected within each journal type (animal behavior,
also included the two highest-impact general science journals, Sci- biodiversity conservation, landscape ecology, and wildlife biology),
ence and Nature, within the multidisciplinary science category in omitting general science journals as they had insufcient included
ISI. urban wildlife articles (n = 2) to compare. For each selected urban
wildlife paper, another literature search was performed on Web
2.2. Publication inclusion/exclusion of Science within the selected publications journal and year pub-
lished. Within the search results, we selected the rst research pa-
Within each target journal, we performed a literature search per that studied wildlife in non-urban areas. If more than one
during AugustNovember 2011 within Web of Science, searching included urban wildlife publication was selected from the same
for the term urban for all publications from 1971 to 2010. The journal and year, additional non-urban wildlife research papers
 notation ensured that terms such as urbanization, subur- were selected for that journal year.
ban, exurban and semi-urban were also detected. Each publi-
cation that resulted from that search was then reviewed to 2.5. Analysis
determine if it met the criteria for inclusion. Publications were in-
cluded only if they represented original research on wildlife (non- We summarized the number of urban wildlife publications in-
domestic animals) located in human-dominated, non-agricultural cluded by our process for each journal and eld (animal behavior,
areas. Review papers, editorials, letters, comments, and book re- conservation, ecology, general science, landscape ecology, wildlife
views were excluded. Reasons for exclusion of reviewed publica- management). We also summarized each as a proportion of the
tions were recorded (Roberts et al., 2006). number of publications by that journal, both overall and by decade.
Similarly, we summarized the proportion of included urban wild-
2.3. Publication categorization life papers with various author afliations, continents on which re-
search was conducted, focal taxa, scientic topics of the studies,
For included publications, we collected data on afliation of and type of study site. We also summarized non-urban papers used
rst author (categories: academic, government, non-governmental for comparison in the same manner. We used chi-squared tests to
organization, and private industry), continent in which research evaluate differences between groups, including testing changes in:
was conducted, and type of taxa studied (mammal, bird, sh, (1) publication rate, author afliation, taxonomic focus, scientic
herpetile, arthropod, and other). We also assessed primary topic, and research site type between decades; (2) author aflia-
research topics of the publication, including: (1) animal behavior: tion, geographic location, taxonomic focus, and scientic topic be-
behavior of individual animals, categorized by topics such as tween urban and non-urban publications; and (3) taxonomic focus
dispersal, home range, habitat selection, mating, nesting, and 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


and Urban Planning (3.7%, 85 publications) and Landscape Ecology
We reviewed 1099 total publications generated by our urban (3.2%, 42 publications). However, Biological Conservation contrib-
keyword search within the 16 target journals from 1971 to 2010, uted the largest volume of publications (117). The wildlife biology
ultimately including 571. Of the 528 excluded papers, 300 journals published the highest percentage of urban wildlife publi-
(56.8%) did not research wildlife, 165 (31.2%) were not research pa- cations during the 1990s but were surpassed in the early 2000s by
pers, and 95 (18.0%) did not occur in urban, suburban, or exurban the landscape ecology journals (Fig. 2). Publication rates signi-
areas. Note that some papers were excluded on multiple grounds. 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,
3.1. Annual trends p > 0.18, df = 3). Qualitative trends, however, suggest slow, steady
increases in publication rates in conservation and wildlife journals,
Rates of publication of urban wildlife literature are increasing but consistently low rates in animal behavior and ecology journals
(Fig. 1). Before 1991, only 14 papers were included as a result of (Fig. 2). General science journals were excluded from the analyses
our process (1 in 19711980 and 13 in 19811990), representing of temporal trends due to exceedingly low number of urban wild-
0.01% of the publications by our target journals in that period. In life papers (n = 2; Table 1, Fig. 2).
19912000, 128 papers studied wildlife in urban areas, represent-
ing 0.17% of overall publications. By 20012010, the rate of urban 3.3. Author afliation
wildlife publications in selected journals had increased to 0.55%,
representing 429 total studies (Fig. 1). Overall, 73.7% of included urban wildlife publications listed an
academic afliation for their rst author (Table 2). In comparison,
3.2. Journals 14.5% of lead authors were employed by a state, local, or national
government, 10.9% were afliated with non-governmental organi-
Of our target journals, Wildlife Research published the highest zations, and only 0.70% worked for private companies. Rates did
proportion of publications addressing urban wildlife topics (4.1%, 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 40.9% (119/291) studied mammals, as compared to Europe, where
in work conducted by academics and a decrease in that conducted 47.5% (56/118) included birds and only 24.6% (29/118) included
by governmental researchers (Table 2). Insufcient publications mammals, and Australia, with 41.9% bird (44/105) and 52.4% mam-
were included in the 1980s (13) or 1970s (1) to warrant statistical mal (55/105) studies. The proportion of mammal studies signi-
analysis. 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
3.4. Geographic focus low sample size, we did not test differences among continents
for other focal taxa, include additional continents, or include ear-
Most urban wildlife studies (291/571, 51.0%) were conducted in lier decades, though a high proportion (53.8%) of the 13 urban
North America, followed by Europe (118/571, 20.7%) and Australia wildlife papers published from 1981 to 1990 focused on mammals
(105/571, 18.4%). Only 6.8% (39/571) of studies occurred in Asia, (Table 3).
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, 3.6. Scientic topic
1.6%).
The most frequently represented scientic topics in urban wild-
3.5. Taxonomic focus life research were animal behavior, conservation, and landscape
ecology, followed by wildlife management and population ecology
The taxa most frequently studied in urban wildlife publications (Fig. 3). Studies focusing on community ecology (9.3%, 53/571), hu-
were birds and mammals, (Table 3), though they were rarely stud- man dimensions (8.1%, 46/571), human-wildlife conict (7.2%, 41/
ied simultaneously (20/571, 3.5%). Arthropods, herpetiles, and sh 571), disease (2.6%, 15/571), and genetics (2.4%, 14/571) were less
were studied less frequently. There were no major shifts in taxa common. The most common topic combination was animal behav-
studied over time; percentages of studies focusing on birds and ior and conservation (42.0%, 240/571), followed by landscape ecol-
mammals were not signicantly different between the 1990s and ogy and conservation (31.9%, 182/571), animal behavior and
2000s (both v2 < 0.19, p > 0.65, df = 1); other taxa and decades landscape ecology (31.0%, 177/571), animal behavior, landscape
had insufcient data to test for changes. In North America, 40.2% ecology, and conservation (26.1%, 149/571), animal behavior and
of urban wildlife publications (117/291) included birds while 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 3.8. Comparing urban and non-urban wildlife publications
management (10.5%, 60/571). Other combinations were repre-
sented in less than 10% of publications. The proportion of included studies conducted on urban wildlife
Landscape ecology research increased signicantly from the compared to that conducted on wildlife in non-urban systems did
1990s to the 2000s (Fig. 3, v2 = 6.82, p = 0.009, df = 1) and there not signicantly differ for any author afliation (all v2 < 0.80,
was a trend towards a concurrent decrease in animal behavior re- p > 0.35, df = 1)) or continent (all v2 < 1.32, p > 0.25, df = 1) except
search (v2 = 2.89, p = 0.09, df = 1); other topics did not signicantly Africa, where there was a slight trend towards a smaller proportion
differ between decades (all v2 < 0.90, p > 0.34, df = 1). In general, of urban (2.8%, 16/571) compared to non-urban (8.7%, 13/150)
the proportion of research conducted on different taxa, and on dif- wildlife studies (v2 = 2.98, p = 0.083, df = 1). Studies on birds also
ferent continents, were similar across topic areas (Figs. 4 and 5), tended to be more common in urban (42.6%, 243/571) compared
though a relatively high proportion of urban wildlife management to non-urban (28.7%, 43/150) areas (v2 = 2.75, p = 0.097, df = 1);
publications focused on mammals (51.8%, 86/166) and were con- the proportion of urban and non-urban wildlife studies did not dif-
ducted in North America (62.7%, 104/166). Population ecology fer for any other taxon (all v2 < 2.36, p > 0.12, df = 1). Finally, com-
studies of urban wildlife were similarly frequent in North America munity ecology topics were less frequent in urban (9.3%, 53/571)
(61.3%, 73/119). 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).
3.7. Research site types

The most frequent study site type within our included publica- 4. Discussion
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- The rate of publication of scientic studies of urban wildlife in-
ies conducted in urban areas, 130 (27.4%) also included research in creased more than tenfold in our target journals since 1971, from
natural habitat. Percentages of urban wildlife publications that fo- less than 0.1% of published articles previous to 1990, to nearly
cused on urban, suburban, or exurban areas were not signicantly 2% between 2001 and 2010. While only 14 urban wildlife publica-
different between the 1990s and 2000s (all v2 < 1.07, p > 0.30, df = tions were published before 1991 in the journals we reviewed, we
1). identied over 400 such studies published between 2000 and
S.B. Magle et al. / Biological Conservation 155 (2012) 2332 29

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, journals, and one of those, Landscape and Urban Planning, clearly fo-
both in terms of number of articles published and in the proportion cuses on urban areas. Wildlife biology and then conservation jour-
of published work that focuses on urban areas. Similarly, in a pre- nals published the next highest percentage of their articles on
vious review of two wildlife biology journals (Journal of Wildlife urban wildlife, with the greatest absolute volume of such studies
Management and Wildlife Society Bulletin), Adams (2005) found that published in Biological Conservation. Miller and Hobbs (2002) noted
the proportion of studies on urban wildlife increased from 12% in that fewer than 6% of the publications published in the journal
the early 1980s to 34% in the late 1990s and early 2000s (Adams, Conservation Biology between 1995 and 1999 included work in ur-
2005). This developing focus likely reects the identication of an ban, suburban, or exurban areas; our own more exhaustive review
important gap in ecological research (Adams et al., 1987; Chace in that journal suggests lower rates (<2% overall) for wildlife stud-
and Walsh, 2006; Marzluff et al., 2008) as well as mounting recog- ies specically. In a similar review, Collins et al. (2000) found that
nition of the rapid expansion of urban areas coupled with an over- only 0.4% of papers published in 9 leading ecological journals be-
all decrease of available alternate habitat for wildlife (Dearborn tween 1995 and 2000 focused on urban species (Collins et al.,
and Kark, 2010; McKinney, 2006). Indeed, we note the emergence 2000). Excluding landscape ecology, we also recorded very low
of a suite of urban ecology journals that publish wildlife studies rates of publication of urban wildlife research in our three target
(e.g., Cities and the Environment, Landscape and Urban Planning, Ur- ecological journals (0.2% overall), providing no evidence for appar-
ban Ecology, Urban Ecosystems), as well as journals that handle hu- ent increase since 2000. Our literature review demonstrated that
man-wildlife conict papers specically, including those in urban such studies were rare in animal behavior journals as well, and
areas (e.g., Human-Wildlife Conicts, Human Dimensions in Wildlife). were exceedingly uncommon in the high-impact general science
These specic journals failed to meet our selection criteria of high- journals. Virtually all reviews on this topic conclude that an in-
impact journals published before 1990 and thus were not included creased focus on urban systems is needed among ecologists (e.g.,
in our analyses, but nonetheless represent important additional Adams, 2005; Chace and Walsh, 2006; Collins et al., 2000; Gehrt
outlets for urban wildlife research. et al., 2010; Mcdonald et al., 2008; Marzluff et al., 2008; Miller
While the trend is increasing, we emphasize that overall rates of and Hobbs, 2002; Pickett et al., 2008). Although our results do indi-
publication of urban wildlife research remain low (<2% in our tar- cate some progress in this direction, such publications are uneven
get journals), particularly considering that urban sprawl and its among disciplines and remain relatively scarce in our target
impacts on biodiversity are accelerating globally. Indeed, urbaniza- journals.
tion is a dominant demographic trend (Marzluff et al., 2008) and The majority (ca. 75%) of published urban wildlife research has
the planet is undergoing an unprecedented urban transition been directed by academics, as assessed by rst-authorship of
(Ramalho and Hobbs, 2012); for the rst time in human history, resulting publications. While studies led by government research-
the majority of humans on the planet live in urban areas, with ers represent the next most common type of urban wildlife study,
nearly all population growth this century expected to be urban it is far less common (ca. 15%). Our comparisons across decades
(United Nations Population Fund, 2007; United Nations, 2010). suggests that this disparity, which was noted in the 1980s (Adams
Urbanization has become a primary agent of habitat loss and frag- et al., 1987) and also is evident between the wildlife management
mentation, and as a result, is also a leading cause of species endan- and conservation elds (Jensen and Krausman, 1993), is not dimin-
germent globally (Czech et al., 2000; Mcdonald et al., 2008). ishing and may even be widening. Our results also suggest that this
Biologists have not emphasized urban study systems for a variety disparity is similar for both urban and non-urban wildlife studies.
of reasons, including a traditional preference to focus on native As publications represent the primary currency for advancement in
species in relatively remote, pristine, and undisturbed ecosystems, academic careers, it is logical to expect academics to publish more
as well as logistical, political, and nancial obstacles to urban re- frequently than other researchers in this eld. However, linking ba-
search (Miller and Hobbs, 2002). sic knowledge to informed practice has been identied as one of
Not only were urban wildlife studies relatively infrequent, but the greatest challenges to the discipline of urban ecology (Marzluff
they also varied considerably among disciplines. The landscape et al., 2008). Moreover, one of the stated goals of urban wildlife re-
ecology journals published urban wildlife research in the highest search is to include non-academic researchers in the production of
proportion, though it should be noted we included only two such 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 mans such as pollination and biological control (Losey and Vaugh-
research and hence may provide novel or unique insights. Our an, 2006); additional study is necessary to fully understand how
study design focused on primary authors, and as such it is possible these functions are altered in urban landscapes and the implica-
that non-academic researchers are becoming more involved in ur- tions of these alterations. Less is known about sh responses to
ban wildlife research, either as non-primary authors or at a level urbanization compared to other taxa, but available studies suggest
typically recognized in acknowledgments. Young and Wolf that native sh abundance and diversity tend to decline, and inva-
(2006) identied a trend towards more non-academic research in sive species increase, in more urbanized areas (Paul and Meyer,
the purely urban journals Urban Ecology and Urban Ecosystems be- 2001; Wolter, 2008). Finally, studies investigating multiple taxa,
tween the periods 19751986 and 19972004, which is encourag- and even more than simply one species, were uncommon, as evi-
ing, but our results reveal little evidence for such a trend for urban denced by the low proportion (9.3%) of studies investigating com-
wildlife publications in the primary animal behavior, conservation, munity ecology. Wildlife studies incorporating multiple species
and ecology literature. and community dynamics were more common outside of urban
In terms of geographic focus, more than half (51.0%) of the ur- systems, and these could help guide similar studies in metropoli-
ban wildlife studies we reviewed were conducted in North Amer- tan areas. Assessing interactions between organisms will be neces-
ica, a proportion similar to wildlife studies conducted in non- sary to better understand ecological dynamics of urban systems
urban areas. Certainly it should be noted that all of our selected (e.g. Crooks and Soul, 1999; Faeth et al., 2005; Crooks et al., 2010).
journals were English-language journals, and most are themselves Urban ecology is an inherently interdisciplinary eld (Marzluff
based in North America, and this may impact our results. However, et al., 2008), and this was reected in the diversity of research top-
all included journals accept international work, and are among the ics addressed in urban wildlife studies. The most common topic for
leading journals in their eld, so we believe this result reects a publications in our target journals was animal behavior, with the
true focus in current research. The relatively large number of stud- majority of these studies focusing on habitat requirements, move-
ies in Europe (20.6%) is expected given that Europe was the rst ment, or dispersal within urban settings. Many studies have inves-
continent to begin urban ecological surveys (Adams, 2005). The tigated behavioral adaptations of wildlife to urban areas, including
substantial representation of Australia (18.5% of studies) is likely changes to activity levels, foraging behavior, and reproduction
partially a result of Australias recent focus on urban wildlife ecol- (Ditchkoff et al., 2006). However, despite the prevalence of studies
ogy (Adams, 2005), but perhaps also was inuenced by our inclu- assessing the behavior of urban wildlife, the animal behavior jour-
sion of Wildlife Research, an international journal based in nals generally published a smaller proportion of urban studies than
Australia. When that journal is omitted, however, the proportion the conservation or wildlife journals, suggesting that the wider
of urban wildlife studies conducted in Australia drops to 9.3%, community of animal behavior scientists, many motivated by the-
which still exceeds all other continents but North America and Eur- oretical and not applied questions (Angeloni et al., 2008), are not
ope. The small number of studies published from Asia, South Amer- yet fully engaged in urban research. Studies regarding the conser-
ica, and Africa are particularly worrisome given that these areas vation of wildlife in urban areas were next most common. Conserv-
are experiencing rapid human population growth and resulting ing species in areas of high human density can be controversial, as
urbanization (Chace and Walsh, 2006) and contain numerous bio- some of these urban species may be invasive, harbor disease, and
diversity hotspots (Myers et al., 2000; Mcdonald et al., 2008); in conict with humans (Angermeier, 1994; Forman et al., 2003; Ur-
Africa specically, wildlife studies tended to be less common in ur- banek et al., 2011). Nevertheless, conservation of urban wildlife
ban compared to non-urban systems. For some studies, a language has numerous benets, including protecting rare and threatened
barrier may have prevented researchers in these areas from pub- species, understanding wildlife response to environmental change
lishing in the top journals, and funding restrictions may have lim- such as global warming, and providing ecosystem services and
ited the scope of such studies to the point where publication in education to humans (Dearborn and Kark, 2010). The relatively
high-impact journals was challenging. Finally, comparative studies new discipline of landscape ecology also has been increasingly ap-
on more than one continent are rare. While logistically difcult, plied to urban wildlife (see also Adams, 2005), partly because the
such research will likely be necessary for discovery and renement large spatial extent of urban areas are directly applicable to
of global urban ecological principles. broad-scale landscape ecology approaches. Moreover, landscape
Birds were the most frequently studied taxon, consistent with ecology also explicitly recognizes landscape context and inter-con-
prior reviews (Adams, 2005; Marzluff et al., 2001), and were also nectedness of systems (Wu, 2008), which can be powerfully ap-
more frequently studied in urban compared to non-urban areas, plied to urban areas that are characterized by complex ows
though mammals were also frequent targets for urban ecological between ecological, socio-economic, and physical processes. Stud-
research, particularly in North America and Australia. Taxonomic ies on the population ecology of urban species previously have
focus did not appear to differ between the 1990s and 2000s, been identied as sparse in wildlife biology journals (Adams,
although of the few urban wildlife papers before 1990, the majority 2005), but a relatively large proportion of our studies (20.9%), as-
focused on mammals. Herpetiles, sh, and arthropods were infre- sessed in a broader range of journals, investigated this topic. Less
quent focal taxa of urban wildlife studies and thus represent taxo- represented were studies of genetics, disease, or human-wildlife
nomic gaps that warrant increased attention in future research conict such as animal-vehicle collisions, landscape damage, or
efforts (see also McIntyre, 2000; Paul and Meyer, 2001; Shochat pet predation, though this has been cited as a growing eld of re-
et al., 2004). However, it should be noted that some journals used search (Adams, 2005). It is notable that while direct assessments of
in this analysis (i.e. Journal of Wildlife Management, Wildlife Society wildlife harm to humans and property were rare, studies of man-
Bulletin) deliberately exclude manuscripts with invertebrates and agement of wildlife, sometimes with the stated goal of reducing
sh as main subjects, thus limiting the appearance of these taxa. these conicts, were much more common. This disconnect may
Amphibians (Hamer and Parris, 2011) and aquatic invertebrates suggest that the degree of damage caused by wildlife should be di-
(Paul and Meyer, 2001) are particularly susceptible to urban im- rectly assessed more often to determine if management action is
pacts, including pollutants originating in urban areas, and may rep- warranted.
resent excellent focal species for studies of ecotoxicology or urban In our review, we included studies conducted across a range of
adaptation. Arthropods also represent the majority of wildlife spe- residential land use types, including urban, suburban, and exurban
cies on Earth (Wilson, 1992) and are critical for ecosystem function systems. Urban systems, characterized by high density develop-
(Bolger et al., 2000), including providing important services for hu- ment, dominated the urban wildlife literature (ca. 83% of studies),
S.B. Magle et al. / Biological Conservation 155 (2012) 2332 31

as would be expected given our focus on urban studies. Suburban continue to advance. At present, however, such advancement re-
areas, land use dominated by lower density residential develop- mains uneven, with important research gaps to be resolved. Urban
ment, are widespread and characterized by generalist and invasive wildlife research is still a young eld with unfullled potential and
species and growing conict between humans and wildlife numerous opportunities for a wide range of scientists from a diver-
(DeStefano and DeGraff, 2003). Nonetheless, suburban wildlife sity of disciplines. We contend that the interdisciplinary science of
studies were relatively uncommon (ca. 15%) and we did not detect understanding the animal species who share our cities has tremen-
any apparent increase in their frequency over time. Exurban re- dous and increasing importance for our urbanizing planet.
search in residential development beyond the urban fringe was
rare, perhaps partly because such regions have been identied as Acknowledgments
important for study only recently (Theobald, 2001). However,
these rural residential areas are critical to ecology and conserva- We thank L. Lehrer for assistance with reviews of publications.
tion because they are frequently located adjacent to protected nat- We also thank S. Gehrt and an anonymous reviewer for helpful
ural areas and are expanding even more rapidly than urban areas comments on the manuscript. Funding for VMH, SBM, and MV
in many regions (Theobald, 2005). Additional wildlife research in was provided by the Lincoln Park Zoo and the Davee Foundation.
suburban and exurban areas therefore would be valuable.
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