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COMMENTARY
Loved or loathed, feral pigeons as subjects in ecological
and social research
Stella Capoccia,
1,
* Callie Boyle
1
and Tedd Darnell
2
1
Department of Biological Sciences, Montana Tech of the University of Montana, 1300 W. Park Street, Butte,
MT 59701, USA and
2
Department of Biology, University of North Dakota, 10 Cornell Street Stop 9019, Grand
Forks, ND 58202, USA
*
Corresponding author. E-mail: scapoccia@mtech.edu
Submitted: 28 June 2018; Received (in revised form): 16 October 2018; Accepted: 19 October 2018
Abstract
The feral pigeon (Columba livia) carries with it a reputation that runs counter to conservation: it is feral, exotic and invasive and
even considered down right filthy. But upon closer inspection, the pigeon could be a subject worthy of study for both ecological
and social science purposes and highlight the importance of urban species in ecology. This insight on pigeons occurred during
an urban ecology study that focused on habitat and population dynamics of the birds and is also supported through an exten-
sive literature review. As a nearly ubiquitous species, we discuss how important pigeons are as a prey-base for numerous rap-
tors and underscore our position by highlighting several studies on the biological uptake of environmental contaminants that
may have benefited from including the pigeon as part of the research. We further our stance on the importance of pigeons in ur-
ban ecology by highlighting additional concerns such as zoonotic disease and climate change. We expand our case by turning to
the social construction of nature, the importance of public participation in conservation. Once again, the ubiquitous presence of
pigeons lends itself well to citizen science in ecology and conservation, especially to show trends across a range of geographic
locations. In short, this commentary strives to reconceptualize the feral pigeon, promoting the bird as a valuable asset to ecolog-
ical and social research in ways that raise awareness for conservation concerns and advance our scientific thinking.
Key words: feral pigeon, urban ecology, keystone species, invasive species, ecological indicators, citizen science
Introduction
This essay promotes the feral pigeon (Columba livia)asan
important subject for urban ecological and social science research.
This position came about during an urban ecology study on
pigeons in Butte, Montana. During our work, we realized the re-
markable place pigeons have in the urban environment.
We suggest that a change in attitude toward and an increase in sci-
entific focus on pigeons could enhance the field of urban ecology
in a variety of ways. Throughout this essay, we focus on the role of
the feral pigeon as a missing link in ecological studies specifically
in urban environments, as a global and local environmental indi-
cator, and as a valuable subject for citizen-science-based projects.
Non-native species, especially non-native birds, are often
understudied (Seymour 2013). Similarly, species of abundance
are often overlooked in research, likely because of their com-
monality, and underfunded, likely because of a lack of urgency
(Hamilton 2008). These two conditions place the pigeon, along
with a few other avian species, as nearly invisible in ecological
research, other than in the context of ecological problems and
population reduction. Yet the pigeon is a global species that
V
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1
Journal of Urban Ecology, 2018, 1–6
doi: 10.1093/jue/juy024
Commentary
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carries with it a wealth of information that can be studied at the
local level. Audubon magazine’s March/April (2013) edition, en-
titled: ‘Why Birds Matter’; elaborates by highlighting topics such
as understanding animals’ social behavior, avian habitat
changes and using birds as both disease indicators and indices
of environmental health. The magazine points to pigeons as a
species that can help detect poor air quality (Berger 2013; see
also: Schilderman et al. 1997), supporting the concept that the
birds may serve as reliable environmental indicators (Morrison
1986). Our position occurs at the intersection of these asser-
tions: the feral pigeon is a surprisingly understudied animal
ecologically but one that holds great potential for providing in-
formation about local and global urban environments.
Compared with other well-populated invasive species, such as
the House Sparrow (Passer domesticus) and European Starling
(Sturnus vulgaris), pigeons are easier to identify as a species, as
individuals, from a distance and track in their home range.
Of equal importance is the role that human interest plays in
conservation, a key component in the urban ecology discourse.
People’s value of nature, how they expect an environment to be
managed, and how people effect or are affected by management
policies, increasingly relies on citizen science research. Steiner
(2016) discusses the importance of stewardship and points to
the role of understanding as critical components in sound plan-
ning. A budding argument exists for using species such as the
pigeon and house sparrows (Passer domesticus) in ecological
stewardship and citizen-science-based activities (Dunn et al.
2006;Larson, Cooper, and Hauber 2016). We build on this
position to promote using the pigeon to help people connect to
urban stewardship in both ecological and social frameworks.
In many ways, the position to advocate for the pigeon is an
uncomfortable one. Naturalists and environmentalists alike es-
pouse sentiments for the bird that are unfavorable at best. The
pigeon carries with it all that runs counter to ecology and con-
servation: these birds are feral, exotic and invasive. The birds
are associated with habitat loss for native species and social
concerns such as disease and filth. Named ‘rats of the sky’,
pigeons are a flagship species of the human imprint on local
biodiversity (Jerolmack 2008). It seems no wonder that pigeons
are underrepresented in studies that examine their positive role
in urban ecology. That said, much of the ecology and social rep-
utation of pigeons stems from a fascinating history intricately
tied to human development and gives credence to our position.
The wild type, or founder species to the common, urban or fe-
ral pigeon, is the rock pigeon of Eurasia, also: C. livia. The natural
habitat of these birds is mixed grasslands for ground foraging
and rocky cliff faces for nesting. The birds were domesticated
over time, selectively bred for high, year-round egg production,
meat and a strong attachment to their territories which aided in
human communication via carrying messages back to their nest-
ing sites (see Blechman 2006; Jerolmack 2013). As with most ani-
mal domestication, the individuals with the highest tolerance for
humans had the greatest access to food which, in turn, led to re-
productive advantages and a growth in populations (see Caras
1996;Trut 1999). So valuable were pigeons to humans that they
were carried across the Atlantic Ocean to the New World
(Blechman 2006; Jerolmack 2013). Today, feral pigeons live pri-
marily in the urban environment around the world, feast largely
on human food waste and can reproduce year-round.
The pigeon’s remarkable ability to live in human environments,
especially the cityscape, is why we consider them a key species in
urban ecology. A species that is both ubiquitous and abundant, the
pigeon is a constant in nearly all urban environments. The pigeon’s
cosmopolitan standing should give them a prominent status in
ecological and social science research as it avails them to compara-
tive studies in different geographic locations. We point to a body of
work that we argue, could be improved upon with the inclusion of
pigeons as a key component to understand certain ecological
issues, such as environmental contaminants. This essay frames
the pigeon as an important species for ecological research, society
and the changing face of environmental conditions.
Ecological import
We began our research in Montana in response to Butte-Silver
Bow County’s request to learn more about the feral pigeons re-
siding in the central business district (CBD), ultimately as a
means to initiate a management program. Our primary research
focused on identifying and characterizing feral pigeon’s nest,
roost and forage sites within the CBD and surrounding neigh-
borhoods. We mapped the distribution of the birds, and, not
surprisingly, found a strong connection between pigeons and
specific human environments: vacant buildings, areas that had
moderate protection from the elements, such as three-side
wind breaks and overhead cover, and a steady food source, like
restaurant waste (Capoccia, Boyle, and Darnell 2014). We also
examined movement of individual pigeons between colonies,
dispersal of juveniles from colonies and estimates of population
size. While simple, this work provides baseline data for the pop-
ulation at hand and allows us to test different research techni-
ques in terms of trapping, marking and re-sighting methods
(Richardson, Capoccia, and Hart 2016).
One of our most unexpected findings was that the pigeon
offers untapped research opportunities. Though not a true wildlife
species, compared with other free-roaming animals, pigeons are
often close at hand, easy to access and capture, and require mini-
mal permits to handle. There are over 20 classifications of plum-
age variants which make visual identification particularly easy
(Dunmore 1968). The pigeon provides opportunities to learn and
experiment with a range of methods pertaining to population dy-
namics, animal behavior, ecological interactions and basic study
design. Additionally, these birds are abundant and universal and,
with increasing urbanization, we argue that it is important to ob-
serve the dynamics of a species that boasts such remarkable con-
sistency in a range of geographies. Studying and tracking pigeon
populations have the potential to elucidate shifts in environmen-
tal conditions, including environmental contaminants, avian dis-
ease, and climate change, that could provide insight into or
forecast shifts in the regional biodiversity (see Hamilton 2008).
So established and integrated are these birds in urban and
suburban environments that we argue they could be considered
an urban keystone species. Pigeons are found on every conti-
nent except Antarctica; the magnitude of such a presence is ac-
companied by a major role in the way habitat is shaped and
nutrients and minerals are cycled. For example, as a species
that naturally evolved to digest seeds, pigeons are known
agents of seed dispersal (Butcher and Bocco 2009) and provide
connectivity between urban islands. Furthermore, a growing
amount of research exists on the limiting factors present in an
urban environment, such as noise, human activity and cover
type, and these factors shape the diversity of species present in
urban habitats (Luniak 2004;Fisher et al. 2016;Aronson et al.
2017). Not only is the pigeon a species that can thrive in an ur-
ban habitat but it is also contributes to the tropic levels in an ur-
ban ecosystem. They are effective as primary consumers to the
extent that their population can support substantial predation
and are an important food source for many birds of prey.
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Countless publications highlight the pigeon as a key
prey species for urban-dwelling raptors across the globe.
For peregrine falcons (Falco peregrinus), in particular, Rejt (2001)
estimated feral pigeons composed a year-round average of 32%
of the diet in Warsaw, Poland with a peak of over 50 % in the
winter and summer. A similar study in southwest England
showed pigeons and doves made up 47% of the prey-base of
peregrines, pointing to pigeons as a key species in the raptor’s
diet (Derwit and Dixon 2008). In Brazil, pigeons make up 68.5%
of the Peregrine diet in Porto Alegre and a stunning 80.9% in
Santos (Silva 1997). Farther east, pigeons were shown as a fa-
vored species in the Peregrine diet in Patagonia, Ria Deseado,
Argentina (Garcia, Bo, and Yorio 2014). In New York City, the es-
timate of pigeons as part of the peregrine’s diet is thought to
reach peaks of 75% during certain time periods (Angier 1991;
Cade 2016). Arizona reports having one of the strongest pere-
grine populations in the United States and asserts that even fal-
cons that nest and roost away from urbanization come into
cityscapes to feed on pigeons (AZGFD 2016). In addition to pere-
grine falcons, pigeons are reported as a key prey species for
other birds of prey that include, but not limited to: 10% of the
diet of urban common kestrels (Falco tinnunculus)(Yalden 1980),
sharp shinned hawks (Accipter striatus)(Mueller and Berger
1970), coopers hawks (Accipiter cooperii)(Boal and Mannan 1999)
and considered the ‘principle avian prey’ of most European gos-
hawk (Accipiter gentilis) populations (Rutz et al. 2006: 580).
Given the domestic pigeon’s important role as a prey spe-
cies, one might go so far as to argue that this bird filled the
prey-base niche left by the Passenger Pigeon (Ectopistes migrator-
ius) in North America. While these two species clearly differ in
flock size, they share the family, Columbidae, much of the same
forage type, and are known prey for the many of the same pred-
ator species (Halliday 1980). The pigeon we know today in-
creased in presence over the last few hundred years, coming in
the wake of the Passenger Pigeon’s decline and extinction in the
early 1900s, offering the predators an alternative. In fact, evi-
dence points to the fact that the population of Cooper’s hawk
changed behavior to switch prey species from the Passenger
Pigeon to our urban Pigeon (Dunne 2017). In addition, the
Peregrine falcon, though marked by dichlorodiphenyltrichloro-
ethane (DDT), is thought to have endured an earlier and much
larger impact by the loss of its key prey, the Passenger Pigeon
(Bodio 2010). It is worthwhile to speculate what the current ac-
cipiter and falcon population might look like if the Passenger
Pigeon had met its fate without its urban successor.
The focus on feral pigeons as an important prey species could
provide insight to and justification for advanced research on ur-
ban birds of prey, specifically regarding environmental contami-
nants and avian disease. As history illustrated with DDT, the
bioaccumulation of environmental toxins has the potential to re-
sult in severe impacts on predatory bird populations. While the
peregrine falcon is often the ambassador species for recovery,
DDT’s impact was wide sweeping and affected a range of species
from the bald eagle (Haliaeetus leucocephalus) to countless passer-
ines (Henry 1977;Rattner 2009;Elliott et al. 2015). In the past few
decades studies have found raptors with biological accumula-
tions of lead (Burger 1995), polybrominated diphenyl ethers
(PBDE) and polychlorinated biphenyls (PCB) (Park et al. 2009;
Newsome 2010) to name a few. One study in particular examined
raptor carcasses and hypothesized that the environmental con-
taminants were a result of bioaccumulation from the carbon
source, though not cited as the cause of death (Elliott et al. 2015).
The article stresses the fact that persistent organic pollution has
the potential to biomagnify as trophic levels increase and goes
on to detail urban expansion and agriculture as likely sources of
chemicals including, but not limited to, PBDEs, PCBs, and 1,1-
dichloroethylene (Elliott et al. 2015). As granivores that frequent
agricultural areas as a food source, pigeons serve as a species
that could make the link between the carbon source, parentheti-
cally noted as ‘representative of a high-corn diet’ (p. 158), organic
pollutants and upper-level predators.
A second study reports on the best samples to use in pan-
continental trends in environmental contamination of birds of
prey (Espin et al. 2016). The article addresses the challenge of
sampling raptors based on their legal protection status and fo-
cuses on biological tissues, such as: feathers, preen oil, car-
casses and crop-content food remains as alternative sampling
methods with varying degrees of success. The authors cross-list
eight alternative sample types at an impressive 35 locations
across Europe and surrounding regions as means to show the
geographic scope of the issue. While comprehensive, the paper
lacks the attention to the prey base (other than in the context of
scavenging), not the least of which is the pigeon, present in all
of the 35 study regions, including Iceland. In both articles, the
feral pigeon could have been an integral part of the research to
map bioaccumulation of toxins, the results of which could go
on to strengthen raptor-focused conservation agenda.
Feral pigeons also make excellent subjects to study other en-
vironmental scenarios such as avian diseases and climate
change. West Nile Virus (WNV) and avian influenza, are known
to infect pigeons and have the potential to be widespread; track-
ing pigeon populations could be a key mechanism to follow a
pandemic. But even diseases that occur at a regional scale, in-
cluding avian cholera, Escherichia coli (E. coli), histoplasmosis and
Salmonella (ssp.), all of which are documented in pigeons, can
cause substantial outbreaks and move between species (Botzler
1991;Panigrahy 1996;Gibbs et al. 2005;Cano-Terriza et al. 2015;
Benedict and Mody 2016). The pigeon’s characteristics of being
easy to find, track and having a year-round presence, lend well
to early identification. Pedersen et al. (2006) tested pigeons in
Fort Collins, Colorado, for E. coli and salmonella. While they did
not detect E. coli, they did determine that 3.2%, of pigeons were
positive for salmonella, which is consistent with the findings of
Toro et al. (1999) from Santiago, Chile. The transmission of any
these diseases between species is not unlikely. Phonaknguen
et al. (2013) pointed out that pigeons and domestic chickens
(Gallus gallus) have the potential to spread the H5N1 strain of
avian influenza when the two species comingle. In a 6-year
study in Brazil, Felippe et al. (2010) showed avian infectious
bronchitis virus also moved between domestic chickens and
pigeons. The same is true for cholera, histoplasmosis, salmo-
nella and E. coli, which brings human health concerns into the
picture.
These and other avian-based diseases can be exacerbated by
fluctuations in environmental conditions such as increases in
temperature and precipitation, factors that are subject to cli-
mate change. It is well-accepted that as environmental stres-
sors increase, body condition tends to decrease which makes
individuals more susceptible to disease. A study in Egypt
showed that Columba livia naturally infected with H5N1 had a
50% mortality rate (Mansour 2014). Another study showed that,
in an outbreak of WNV, while pigeons did not show statistically
significant mortality, they have a 70% infection rate, which
could then translate into high transmission rates (Bakonyi et al.
2013). Monitoring residential bird populations, combined with
gathering data on environmental variables, like weather, is a
great way to detect changes in populations that are often asso-
ciated with the aforementioned diseases.
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Regardless of the scenario, the feral pigeon offers a relatively
consistent variable from which a host of ecological conditions
can be studied. Pigeons are a standard subject that provide con-
nectivity between places such as Beijing, China and Butte,
Montana. The relationship pigeons create among urban envi-
ronments could be considered a touchstone for ecological
health. Literature on abundant species management refer to
both native and exotic species as important populations from
which to monitor general environmental conditions. Moreover,
history reminds us that the decline of abundant species, partic-
ularly prey species, could result in a subsequent trophic cas-
cade, so despite high numbers, the vulnerability of abundant
populations should not be discounted (Hamilton 2008). In short,
major oscillations in pigeon populations are uncommon. As a
result, the bird’s omnipresence makes them an ideal study spe-
cies because they provide consistent data that is less noisy and
easier from which to draw inferences in scientific studies.
Citizen science and a link to conservation
As ecological management expands to include the urban land-
scapeandhumanprioritieswecan,onceagain,turntothepigeon
for research. Local ecology, defined as the area closest the user,
networks into larger ecological processes and informs us about
our own habitat. When it comes to collecting data on the local
and urban ecology, wildlife scientists often benefit from under-
standing the community knowledge of a given species (Cooper
et al. 2007;Belt and Krausman 2012;Haywood, Parrish, and
Dollivert 2016) and understanding the attitudes people have to-
ward the animal species (McCleery et al. 2006). Though the out-
ward view is that pigeons are problem animals linked to
destruction of buildings and potential spread of disease, closer
inspections reveal that it is quite common to find people who
have a strong attachment to the birds, and others who simply feel
pigeons belong in the area (see Jerolmack 2013; Capoccia, Boyle,
and Darnell 2014). In fact, feral pigeons are viewed more similarly
to other free-roaming wild animals than domestic ones and serve
as a valuable link to the natural world, for good or for bad (Dunn
et al. 2006; Jerolmack 2013; Capoccia, Boyle, and Darnell 2014).
People who favor pigeons are passionate about their pres-
ence to the extent that they know individual birds and feed
whole flocks, aspects that contribute to the birds’ ecological car-
rying capacity. Many people know which directions pigeons
come from in the mornings, where they head in the evenings,
and even seasonal differences in these behaviors. In contrast,
those who do not like pigeons remark about other components
of the urban ecology, noting that they often see pigeons comin-
gling with ravens or that they saw a fox or raptor by their house
and hope it is preying on pigeons. In both cases, community
members addressed pigeons as part of nature and show famil-
iarity about behavior, population dynamics and even other spe-
cies (Jerolmack 2013; Capoccia, Boyle, and Darnell 2014). The
pigeon’s integration with nature aligns with work by Dunn et al.
(2006) who state that feral pigeons are a significant connection
between urbanites and the natural world. They go on to support
the idea that the relationship to the natural world is critical for
conservation, a position that is espoused by many scholars. In
fact, Castree and Braun (1998) quote Neil Smith saying, ‘nature
separate from society has no meaning’ (p. 6).
Attitudinal research on nature and conservation stems from
citizen science and conservation. In fact, the discourse on human
dimensions of natural resource management stresses the impor-
tance of understanding human attitudes and public involvement
as they relate to wildlife, conservation and ecological
management. Nowhere could this be more important than on the
urban landscape where the general public become the stakehold-
ers. People’s investment in pigeons shows relationships to nature
and animals, where on the contrary, negative attitudes tend to re-
flect the degradation of the environment (Jerolmack 2013).
McCleery et al. (2006) point out that people’s attitudes toward
wildlife forecast human behaviors and that human value brings
about motivation to support conservation efforts for a range of
species.
For conservation to be a success we require widespread sup-
port of and participation in environmental stewardship. Calling
upon the local community to track and record animal sightings
is an excellent way to engage the public in science (Larson,
Cooper, and Hauber 2016;Rudd et al. 2016). And, when it comes
to collecting data on the local ecology, wildlife scientists often
benefit from understanding the local knowledge of a given spe-
cies and, in turn, community involvement improves steward-
ship (Belt and Krausman 2012;Haywood, Parrish, and Dollivert
2016). Larson, Cooper, and Hauber (2016) show the importance
of citizen science monitoring house sparrows. They point out
that understanding people’s emotions provides a link to how
they value wildlife which can contribute to local ecology regula-
tions. A complimentary study shows the importance of commu-
nity perspectives in designing management plans for white-
tailed deer in Pennsylvania. Improved communication between
the general public and the wildlife specialists improved the sat-
isfaction with the deer management plan (Fleegle, Rosenberry,
and Wallingford 2013). Results from a citizen science project in
California showed that community members excelled at identi-
fying health problems in a local squirrel population. The com-
munity members were able to identify six different clinical
signs of notoedric mange (Notoedres centrifera) in the squirrels
and did so faster than the local officials (Rudd et al. 2016).
Results are similarly impressive from studies on the health and
population dynamics of mountain goats (Oreamnos americanus)
in Glacier National Park, Montana, USA (Belt and Krausman
2012) and koala (Phascolarctos cinereus) in Australia (Predavec
et al. 2016). In all cases, the authors report accuracy in species
knowledge and comprehensive geographic coverage, informa-
tion which the scientific team alone was unable to acquire.
Across the field of ecology there is an increasing emphasis
on the importance of communities’ knowledge of population
changes over time and even on individual animal sightings that
may improve accuracy of territory data and population sizes.
Collectively, these examples demonstrate the strength of hu-
man perspectives in ecological management and the impor-
tance of people’s investment in conservation issues. These
finding help scientists and policy makers develop management
plans by gaining a broader understanding of an areas biodiver-
sity, the shifts in animal movement and how the social culture
relates to nature (McCleery et al. 2006;Larson, Cooper, and
Hauber 2016). Understanding a community’s tolerance level for
a species helps inform the management protocol, especially
with regard to population reduction (McCleery et al. 2006;
Fleegle, Rosenberry, and Wallingford 2013;Larson, Cooper, and
Hauber 2016). These studies validate the fact that social per-
spectives and urban ecology are both valuable for conservation
and gaining traction in the arena of wildlife management.
Given the widespread distribution of pigeons, we argue that
community efforts to monitor local pigeon populations would
be an ideal way to link people into larger discussions on topics
such as avian health or climate change. The pigeon could serve
as a catalyst to start the discussion regarding people’s perspec-
tives on the surrounding ecology as well as the nature of
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invasive species. The bird’s robust size, low flight response and
seasonal omnipresence make them an ideal species for people
to connect with. Their unique coloring and strong attachment
to a territory help people identify individual birds and reduce
over counting. And, as stated earlier, pigeons do not require ex-
cessive permitting, they are easy to handle, and can be marked
using plastic colored leg bands made for the poultry industry, so
their population can be tracked with remarkable ease and accu-
racy. The strong human–animal interactions that occur be-
tween pigeons and people could potentially be a staple for
information on urban wildlife populations.
Conclusion
We have in no way exhausted all the avenues for research in-
volving the feral pigeon. Pigeons serve as a reflection of our en-
vironmental impact and can be used to monitor the
environmental condition. This bird is one of several species
that mark human advancement: from early domestication to
our own high-density living. As our environment continues to
flux, a free-roaming and pervasive animal perches right out-
side our front door, sometimes literally, and provides an echo
to our actions. While the pigeon’s presence initially represents
a negative shift in biodiversity as an invasive species, its pres-
ence holds an important role in ecosystem function, not the
least of which is the urban setting. Changes in the pigeon’s
population can be used as an assessment of environmental
conditions, and a way to connect local environments. In many
ways, the pigeon is as urban as people are and may be our
modern canary.
Pigeons have the potential to serve as a gateway species
for scientists to explore the relationship people have with
their local environment, and promote science and conserva-
tion. Well over a century ago, the human attachment to nature
was espoused by George Perkins March, Man and Nature (1864),
followed by Aldo Leopold, A Sand County Almanac (1949) and
later, Roderick Nash, Wilderness and the American Mind (1964).
Today, human interests remain a core part of contemporary
conservation which turns to urban and suburban settings as
emerging sites and citizen science for new methodological
approaches (Decker, Brown, and Siemer 2005). As we continue
to increase our collective understanding of our environment,
we can take research into the future using the pigeon as a
powerful lens to examine the urban dynamics of ecological
needs.
Nature is not always the wild conditions promoted through
places like natural parks, it can also be found in the trees that line
urban streets (Cronon 1996;DeStefano 2010;Dunne 2017). Loved
or loathed, pigeons have the ability to give us a much better un-
derstanding of urban wildlife interactions that could be relevant
to a variety of other wildlife species and has the potential to vindi-
cate the critical role urban ecology has on conservation at large.
Acknowledgements
Tremendous appreciation goes to Richard Douglass, Ed
Randall, Dan Powers, Matt Vincent and Butte-Silver Bow
County for their commitment to this project from the start.
A special note of appreciation goes to those helped edit this
work, especially Amy Kuenzi, Lindsay Torpey, Jay Sellmer
and Cody Richardson. We also thank Montana Tech of the
University of Montana for providing the institutional
support for this manuscript. And to Bill and Asha Henne for
being patient during this work.
Conflict of interest statement: None declared.
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