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Coyote (Canis latrans) diet in an urban environment: variation relative to pet conflicts, housing density, and season

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Coyotes (Canis latrans Say, 1823) are highly successful in urbanized environments, but as they populate cities, conflict can occur and often manifests in the form of incidents with pets. To better understand whether coyotes view pets as prey or, alternatively, as competitors or a threat, we conducted a diet analysis of coyotes in the Denver metropolitan area (DMA) by analyzing scats. We also examined differences in diet between high-and low-density housing and among seasons. We found only small percentages of trash and domestic pets in the coyote diet. The presence of pets in the diet did not coincide with the increase of pet conflicts in the DMA in December and January, supporting the hypothesis that coyote conflict with pets is primarily driven by competition or a threat response. Coyotes relied mostly on native plant and animal species, and rodents and lagomorphs were the most prevalent diet items. Coyotes consumed rodents and non-native plants more often in high-density housing and deer, corn, and native plants more often in low-density housing. Coyotes also consumed more fruits and invertebrates during summer and autumn and more mammals and birds in winter and spring. As human–coyote conflicts increase in urban areas, understanding how coyotes and other urban-adapted carnivores use anthropogenic resources may provide insight that can be used to promote coexistence between humans and wildlife.
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Coyote (Canis latrans) diet in an urban environment: variation
relative to pet conflicts, housing density, and season
S.A. Poessel, E.C. Mock, and S.W. Breck
Abstract: Coyotes (Canis latrans Say, 1823) are highly successful in urbanized environments, but as they populate cities, conflict
can occur and often manifests in the form of incidents with pets. To better understand whether coyotes view pets as prey or,
alternatively, as competitors or a threat, we conducted a diet analysis of coyotes in the Denver metropolitan area (DMA) by
analyzing scats. We also examined differences in diet between high- and low-density housing and among seasons. We found only
small percentages of trash and domestic pets in the coyote diet. The presence of pets in the diet did not coincide with the increase
of pet conflicts in the DMA in December and January, supporting the hypothesis that coyote conflict with pets is primarily driven
by competition or a threat response. Coyotes relied mostly on native plant and animal species, and rodents and lagomorphs were
the most prevalent diet items. Coyotes consumed rodents and non-native plants more often in high-density housing and deer,
corn, and native plants more often in low-density housing. Coyotes also consumed more fruits and invertebrates during summer
and autumn and more mammals and birds in winter and spring. As human–coyote conflicts increase in urban areas, under-
standing how coyotes and other urban-adapted carnivores use anthropogenic resources may provide insight that can be used to
promote coexistence between humans and wildlife.
Key words: anthropogenic, Canis latrans, coyote, Denver, food habits.
Résumé : Les coyotes (Canis latrans Say, 1823) ont beaucoup de succès dans les milieux urbanisés, mais leur établissement dans
les villes peut entraîner des conflits qui se manifestent souvent par des incidents impliquant des animaux de compagnie. Afin
de mieux comprendre si les coyotes considèrent les animaux de compagnie comme des proies ou comme des concurrents ou
menaces, nous avons analysé les régimes alimentaires de coyotes dans la région métropolitaine de Denver (DMA) en analysant
des excréments. Nous avons également examiné les différences de régime alimentaire entre des secteurs résidentiels de forte
et de faible densité et d’une saison a
`l’autre. Nous n’avons décelé que de faibles pourcentages de déchets et d’animaux de
compagnie dans le régime alimentaire des coyotes. La présence d’animaux de compagnie dans l’alimentation des coyotes ne
coïncidait pas avec l’augmentation des conflits avec ces derniers dans la DMA en décembre et janvier, ce qui appuie l’hypothèse
voulant que les conflits entre coyotes et animaux de compagnie découlent principalement de la concurrence ou de réactions
`la menace. Les coyotes avaient principalement recours a
`des espèces de plantes et d’animaux indigènes, les rongeurs et
lagomorphes étant les éléments prédominants de leur régime alimentaire. Les coyotes consommaient plus souvent des rongeurs
et plantes non indigènes dans les secteurs résidentiels de forte densité, et des cerfs, du maïs et des plantes indigènes dans les
secteurs résidentiels de faible densité. Les coyotes consommaient également plus de fruits et d’invertébrés durant l’été et
l’automne et plus de mammifères et d’oiseaux a
`l’hiver et au printemps. Avec l’augmentation du nombre de conflits entre
humains et coyotes dans les zones urbaines, la compréhension de l’utilisation des ressources anthropiques par les coyotes et
d’autres carnivores adaptés au milieu urbain pourrait fournir des renseignements utiles pour les efforts visant a
`promouvoir la
coexistence des humains et des espèces sauvages. [Traduit par la Rédaction]
Mots-clés : anthropique, Canis latrans, coyote, Denver, habitudes alimentaires.
Urbanization can lead to habitat loss and fragmentation and is
one of the primary causes of species endangerment (Czech et al.
2000;McKinney 2002;Markovchick-Nicholls et al. 2008). However,
some wildlife species, including certain mammalian carnivores,
can thrive in urban environments (Ditchkoff et al. 2006;Bateman
and Fleming 2012). Carnivores that adapt to urban environments
generally tend to be small to medium-sized, have high reproduc-
tive potential, can tolerate people, and are dietary generalists
(Fuller et al. 2010). The diet of these urban carnivores usually
includes some anthropogenic food, such as cultivated plants, pets,
garbage, and roadkill (Bateman and Fleming 2012).
Coyotes (Canis latrans Say, 1823) live in nearly every major metro-
politan area in the United States (Poessel et al. 2017), and they
exemplify the characteristics of urban-adapted carnivores (Morey
et al. 2007;Gehrt and Riley 2010). Diet studies of coyotes have been
conducted in several urban areas, revealing that coyotes use both
natural food items (e.g., deer, rabbits, small mammals, and wild
Received 5 February 2016. Accepted 15 January 2017.
S.A. Poessel.* Department of Wildland Resources, Utah State University, Logan, UT 84322, USA.
E.C. Mock.
Department of Fish, Wildlife, and Conservation Biology, Colorado State University, Fort Collins, CO 80521, USA.
S.W. Breck. USDA Wildlife Services, National Wildlife Research Center, Fort Collins, CO 80526, USA.
Corresponding author: S.A. Poessel (email:
*Present address: Forest and Rangeland Ecosystem Science Center, U.S. Geological Survey, Boise, ID 83706, USA.
Present address: Denver Zoological Foundation, Denver, CO 80205, USA.
This work is free of all copyright and may be freely built upon, enhanced, and reused for any lawful purpose without restriction under copyright or database
law. The work is made available under the Creative Commons CC0 1.0 Universal Public Domain Dedication (CC0 1.0).
Can. J. Zool. 95: 287–297 (2017) Published at on 26 February 2017.
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fruits) and anthropogenic foods (e.g., garbage, domestic pets, pet
food, and cultivated plants) (McClure et al. 1995;Quinn 1997;
Fedriani et al. 2001;Morey et al. 2007;Gehrt and Riley 2010;
Lukasik and Alexander 2012). Some coyote diet studies have
recognized temporal fluctuations in diet (e.g., Litvaitis and Shaw
1980;Crimmins et al. 2012), but no studies have evaluated diet
relative to urban characteristics, such as housing density.
Increased coyote presence in more urban areas can lead to a rise in
encounters and conflicts with humans (Baker and Timm 1998;
Curtis and Hadidian 2010;Gehrt and Riley 2010). Reports of coyote
conflicts with humans and especially domestic pets occur throughout
the United States (Poessel et al. 2017). For example, in the Denver
metropolitan area (DMA), conflicts with pets are now common,
and behaviors that include stalking and, in rare cases, attacks on
humans have become more prevalent (Poessel et al. 2013). Con-
flicts have been found to be particularly high in the winter months in
both the DMA and Chicago (Gehrt and Riley 2010;Poessel et al.
2013), although increased conflicts during the summer months
have been reported in other North American urban areas (White
and Gehrt 2009;Lukasik and Alexander 2011). Questions about
why pet conflicts have increased and why they fluctuate season-
ally are common among natural resource managers. One question
is whether coyote–pet conflict is driven primarily by the coyote’s
desire to eat pets (i.e., predation) or whether conflict is primarily
a product of coyotes viewing pets as competitors or as a threat.
The answer to this question should allow wildlife managers to
better understand coyote behavior and motivations in highly
urbanized areas.
The principal goal of our study was to identify patterns in the
anthropogenic portion of the coyote diet in the DMA. Our primary
objectives were to (i) examine how dogs (Canis lupus familiaris L.,
1758) and cats (Felis catus L., 1758) vary seasonally in the coyote diet
and how this variation coincides with seasonal trends in conflicts
and (ii) determine how other anthropogenic food items (e.g.,
trash) in the diet vary between areas of high- and low-density
housing (hereafter, “high-density” and “low-density”). Our sec-
ondary objective was to describe the food habits of coyotes in the
DMA and to determine how coyote diet varied in high- and low-
density areas, as well as across seasons. If coyotes viewed pets
primarily as prey rather than as competitors or a threat, then we
expected pet consumption to be higher in the winter months,
mirroring documented patterns in conflict. We also expected that
consumption of trash would be more apparent in high-density
housing areas, reflecting a greater influence of anthropogenic
food sources in higher density areas.
Materials and methods
Study sites
We defined the DMA as the “Denver urban area” as delineated
by the U.S. Census Bureau (United States Census Bureau 2015). The
DMA comprises over 35 municipalities in north-central Colorado
and all or parts of seven counties (Adams, Arapahoe, Boulder,
Broomfield, Denver, Douglas, and Jefferson). It has a semiarid
climate and, during the study, had annual precipitation of 43 cm
and monthly temperatures ranging from a mean low of −10 °C in
December to a mean high of 31 °C in August (Weather Underground
2015). The DMA had a human population size of almost 2.4 million
in 2010 and an approximate size of 1760 km
. Located along the
front range of the Rocky Mountains of Colorado, the DMA lies
between the foothills of the Rocky Mountains to the west and
agricultural fields and grasslands to the east. Historically, the
DMA was primarily dominated by grassland habitat, but now
incorporates various land-cover types, including urban develop-
ment, woodlands, agriculture, and grasslands (Poessel et al. 2013).
Before selecting our study sites, we first defined areas of high-
and low-density housing within the DMA. Designation of these
areas was based on housing-density data obtained from the Spa-
tially Explicit Regional Growth Model (SERGoM version 3; Theobald
2005), which depicts housing density for the coterminous United
States at 100 m resolution. We defined high-density areas as those
with a housing density of less than 0.5 acre per unit and low-
density areas as those with a housing density of greater than
5 acres per unit. High-density areas contained smaller amounts of
open space with housing developments extending to the borders
of such open space, whereas low-density areas contained larger
amounts of open space with housing developments nearby. We
chose to include only high- and low-density areas (excluding
medium-density areas) to compare differences in coyote diet be-
tween two distinctive types of urban habitat. We then selected
11 parks or open-space areas known to contain coyotes near neigh-
borhoods, 5 in areas of high-density and 6 in areas of low-density
housing (Fig. 1). These open-space areas consisted of natural hab-
itat and contained both paved and dirt trails. Within each open-
space area, we then selected trails that we walked to collect coyote
scats. We selected trails used by coyotes (as indicated by the pres-
ence of coyote scats) that covered most parts of the open-space
Scat collection and identification
We collected coyote scats every month for 1 year, beginning in
June 2013. We initially walked each transect to clear all scat and
then subsequently walked each transect once per month to collect
scat, although we only used scat from every other collection pe-
riod in the analyses. We received heavy snow in February that
made collecting scat difficult, but also degraded scat when the
snow melted, leaving few if any samples to be collected; thus, we
were unable to analyze scat for the month of February. We were
also unable to conduct genetic analyses during our study; hence,
we distinguished coyote scats from dog scats based on color,
shape, texture, and the amount of hair. Coyote scats tend to be
12–30 mm in diameter; red, dark brown, or gray in color; contain
bones, hair, grasses, or seeds; and possess thick, segmented cords
and tapered ends (Lukasik and Alexander 2012). Dog scats can be
many sizes, are brown or yellow in color, tend to be smoother in
shape, and lack hair, bones, and seeds. We removed any scats that
looked similar to dog scats; thus, we possibly may have discarded
coyote scats that may have contained a diet similar to a domesti-
cated dog. We did not observe any foxes or bobcats (Lynx rufus
(Schreber, 1777)) in our study sites (so we assumed they were rare),
and raccoon (Procyon lotor (L., 1758)) scats have a different shape
than coyote scats, so we were not concerned about differentiating
coyote scats with those of these species. We stored scats in a
laboratory at room temperature until analysis.
Scat analysis
We randomly chose up to 10 scats per site for the months of
August, October, December, March, April, and June and combined
these scats into a single fecal sample. If we did not have 10 scats for
a particular site (which frequently occurred), then we used all of
the scats available. We thus had one fecal sample per site per
month for analysis purposes. We chose this analysis method not
only because of limited resources to process each scat individu-
ally, but also because it allowed us to address our primary ques-
tion of whether the importance of anthropogenic items varied
between seasons or between housing densities.
We first brushed off each scat to remove any external vegeta-
tion or rocks that may have attached to the scat after defecation.
After combining scats into fecal samples, we then placed them on
a tray and dried them in an incubator at 50 °C for 24 h. We
recorded the total dry mass and then placed the fecal sample in a
soap bath for 24 h to soften the feces for analysis.
Once softened, we broke up and washed the fecal sample
through a sieve that retained all of the macroscopic material of
hair, feathers, exoskeletons, bones, seeds, vegetation, and trash.
We discarded the microscopic material (more than 50% of the
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fecal sample), containing unidentifiable organic matter but also
potentially earthworm chaetae and feather fragments (Reynolds
and Aebischer 1991). We washed and dried the remaining material
(i.e., macroscopic material) two more times after placing material
in a large bucket with water and mixing it with a paint mixer. Our
goal was to separate bones, seeds, and trash, which tended to sink
from the hair, feathers, vegetation, and exoskeletons that tended
to float. We then separated bones, hair, seeds, and trash and re-
corded the dry mass of each of these items. We then divided the
masses of each of these components by the total mass of the fecal
samples to calculate the percent composition. These percentages
provided a relative index of vertebrates (bones, which could in-
clude mammals, birds, reptiles, and amphibians), mammals (hair),
plants (seeds), and anthropogenic food (trash) in the coyote diet.
We subsampled hairs from each combined fecal sample to iden-
tify the mammals consumed by coyotes. For each fecal sample, we
randomly selected 100 hairs using a system of gridded cells on a
tray. We then laid the selected hairs parallel to one another on a
slide for analysis under a microscope. Using a hair identification
key, we identified each subsampled hair according to banding,
medulla, and cuticle patterns and placed each hair into the following
categories: rodent, lagomorph, deer, raccoon, red fox (Vulpes vulpes
(L., 1758)), coyote, cat, dog, soricomorph, and unidentified (Moore
et al. 1974). We further condensed these categories by combining
Fig. 1. Map of the western portion of the Denver metropolitan area (DMA) with high- and low-density housing. White space includes public
land, undeveloped private land, and areas of housing density between 0.5 and 5 acres per unit. Black crosses represent the five high-density
transects and black stars represent the six low-density transects used to collect coyote (Canis latrans) scats from 2013 to 2014.
Poessel et al. 289
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raccoon, red fox, and coyote into a “carnivores” category and cat
and dog into a “pets” category. We recorded the frequency of each
mammal category in fecal samples and calculated the percentage of
hairs in each mammal category by dividing the number of hairs in
each group by the total hairs analyzed for each sample (i.e., 100 hairs).
We then computed mean percentages for each housing density
across months and for each month across housing densities.
Because we used hairs to identify mammal species (the most
common vertebrate group in fecal samples), we did not identify
the bones in the fecal samples to species or use the mass of bones
in analyses. We identified birds by analyzing feathers (as de-
scribed below). We only found reptile skin in one scat; thus, we
did not include reptiles and amphibians as a separate category in
the coyote diet.
We separated all seeds by type based on size and appearance
and submitted them to the Colorado State Seed Laboratory (Colo-
rado State University, Fort Collins, Colorado, USA) for identifica-
tion. After identification of the seeds to plant species, we then
recorded the masses of seeds by species and the frequency of each
species in fecal samples. We then computed mean seed masses of
each species for each housing density across months and for each
month across housing densities. We further identified each plant
species as either native or non-native.
Finally, in each fecal sample, we visually estimated the amount
of feathers and exoskeletons, which were not removed from the
samples. We did not identify feathers to bird species; however,
exoskeletons were mostly intact and were easily identified as
grasshoppers or beetles. We assigned a separate score for both
feathers and exoskeletons in each sample. For feathers, we as-
signed a 0 when no feathers were visible, 1 when roughly 1%–20%
of the sample was composed of feathers, and 2 when >20% of the
sample was composed of feathers. For exoskeletons, we assigned a
0 to fecal samples with no exoskeletons present, 1 when
roughly 1%–10% of the sample was composed of exoskeletons,
and 2 when >10% of the sample contained exoskeletons. We inad-
vertently used a 10% break for exoskeletons rather than 20%; how-
ever, we proceeded with this scoring rather than simply reporting
presence or absence so that we could present patterns similar to
what we demonstrated for feathers. For both feathers and exo-
skeletons, we then calculated a weighted average of scores for
each housing density across months and for each month across
housing densities. To do so, for each housing density, we multi-
plied the number of sites within a month with a particular score
by the value of that score (i.e., 0, 1, or 2), added these three prod-
ucts, then divided this sum by the total number of sites in a month
for which we had fecal samples. These scores reflect a rough qual-
itative assessment of the mean consumption of birds and inverte-
brates by coyotes (i.e., a score of 0 reflects no consumption, a score
of 1 reflects low consumption, and a score of 2 reflects high con-
We note that, for each food-item group described above (mam-
mals, plants, and birds or invertebrates), we used a different anal-
ysis method to determine variation in the coyote diet between
housing densities and across months. Our goal was to determine
not only how frequently a food item occurred in the diet, but also
how important each item was in the diet by using masses when
possible. These methods may vary from other coyote diet studies.
Statistical analysis
We focused our statistical analysis on our primary objective of
whether coyotes kill pets because of predation or because of com-
petition or a threat response. We conducted an analysis of vari-
ance (ANOVA) test in R version 0.99.446 (R Core Team 2015)to
determine if the amount of pet hair in the coyote diet varied by
Fig. 2. Mean (±SE) percent composition of coyote (Canis latrans) fecal samples (mass of each item divided by mass of the fecal sample) in four
diet groups (bones, seeds, hair, and trash) in high- and low-density sites in the Denver metropolitan area, from 2013 to 2014.
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month. Poessel et al. (2013) previously found that coyote–pet con-
flicts were more frequent during the winter months (specifically,
December and January). If coyotes pursue pets primarily for con-
sumption, then pet hair in scats should increase in December. If
no such pattern occurs, then the hypothesis that coyotes pursue
pets primarily because they perceive them as competitors or a
threat is supported.
We found very little trash in the coyote diet in both housing-
density areas (see Results), so statistical tests to address our second
primary objective, i.e., to determine how anthropogenic food
items vary between high- and low-density housing areas, were not
possible or necessary. We also did not statistically analyze specific
food items and how they varied by housing density or month (i.e.,
our secondary objective). Instead, we present these results in a
Table 1. Frequencies and percentages of mammal hairs, as well as frequencies, masses, and percentages of seeds of
plant species and trash, found in 64 coyote (Canis latrans) fecal samples (consisting of up to 10 scats per site per month)
in the Denver metropolitan area, from 2013 to 2014.
Group Species Frequency Mass (g) Percentage (%)
Rodents NA 64 43.02
Lagomorphs NA 61 25.73
Deer Mule deer, Odocoileus hemionus 35 10.29
Carnivores Raccoon, Procyon lotor 40 8.98
Carnivores Red fox, Vulpes vulpes 34 5.14
Carnivores Coyote 23 1.58
Pets Domestic cat, Felis catus 24 2.12
Pets Domestic dog, Canis lupus familiaris 25 1.21
Soricomorphs NA 10 1.39
Unknown NA 14 0.54
Total by group
Rodents 64 43.02
Lagomorphs 61 25.73
Deer 35 10.29
Carnivores 56 15.70
Pets 41 3.33
Soricomorphs 10 1.39
Unknown 14 0.54
Non-native Russian olive, Elaeagnus angustifolia 17 30.05 6.48
Native Cactus, Opuntia spp. 11 209.28 45.16
Non-native Corn, Zea mays 8 24.47 5.28
Non-native Field bindweed, Convolvulus arvensis 7 0.32 0.07
Non-native Sorghum grain, Sorghum spp. 6 1.44 0.31
Native American plum, Prunus americana 6 24.11 5.20
Native Sunflower, Helianthus annuus 6 0.28 0.06
Non-native Grapevine, Vitis spp. 5 16.60 3.58
Native Manzanita/bearberry, genus Arctostaphylos Adans. 5 2.89 0.62
Non-native Wheat, genus Triticum L. 5 0.40 0.09
Native Chokecherry, Prunus virginiana 4 139.96 30.20
Non-native Watermelon, Citrullus lanatus (Thunb.) Matsum. & Nakai 3 0.16 0.03
Non-native Oat, Avena sativa L. 3 1.96 0.42
Native Virginia groundcherry, Physalis virginiana Mill. 2 0.45 0.10
Native Raspberry, genus Rubus L. 2 0.08 0.02
Native Flax, Linum usitatissimum L. 2 1.42 0.31
Non-native Safflower, Carthamus tinctorius L. 1 0.02 0.01
Non-native Dandelion, genus Taraxacum F.H. Wigg 1 0.06 0.01
Non-native Pigweed, genus Amaranthus L. 1 0.01 0.00
Non-native Black bindweed, Fallopia convolvulus (L.) Á. Löve 1 0.47 0.10
Non-native Hairy vetch, Vicia villosa Roth 1 0.02 0.01
Non-native Alfalfa, Medicago sativa L. 1 3.93 0.85
Unknown NA 19 5.05 1.09
Total mass 463.43 100.00
Total by group
Native 27 378.47 81.67
Non-native 35 79.91 17.24
Unknown 19 5.05 1.09
NA NA 27 21.0 0.31
Note: Group represents condensed categories for mammals and species origin for plants. Frequency represents the number of fecal
samples (out of 64) containing that food item. Mass represents the total mass of each plant species and group and of all trash found in
fecal samples. For hair, percentage is the mean percentage of hairs for each mammal species or group, calculated as the number of
hairs for that species or group divided by total hairs analyzed (i.e., 100) in each fecal sample. For seeds, percentage is the mass for each
plant species or group divided by the total mass for all plant species (463.43 g). For trash, percentage is the mass of all trash divided by
the total mass of all fecal samples analyzed (6788.66 g). NA, not available or not identified.
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descriptive manner by graphically evaluating patterns in the data;
the error bars in these graphs indicate the amount of variation
between housing densities and across months.
Across our 11 sites, we sampled 26.2 km of trails (high-density
housing: 12.5 km, mean ± SD = 2.5 ± 1.4 km; low-density housing:
13.7 km, mean ± SD = 2.3 ± 1.3 km). We analyzed 64 fecal samples,
29 in high-density sites and 35 in low-density sites. These fecal
samples consisted of 424 scats, 210 in high-density sites (mean ±
SD = 7.2 ± 3.0) and 214 in low-density sites (mean ± SD = 6.1 ± 2.9).
Among diet items, hair had the highest percentage by mass in
combined coyote fecal samples (mean ± SE = 26.2% ± 1.7%), fol-
lowed by bones (10.4% ± 1.0%) and seeds (5.7% ± 1.5%). Percentage of
trash by mass in fecal samples was negligible (0.3% ± 0.1%). Hair
percentages by mass were higher in fecal samples in high-density
sites and seed percentages were higher in low-density sites (Fig. 2).
Based on the hair analysis, the most common mammal group
found in fecal samples was rodents (43.0% ± 3.0%), followed by
lagomorphs (25.7% ± 2.7%) and carnivores (15.7% ± 1.8%; raccoon:
9.0% ± 1.5%; fox: 5.1% ± 1.0%; coyote: 1.6% ± 0.3%; Table 1). Carnivore
consumption could have been from scavenging, predation, or
in the case of coyotes, self-grooming. Fecal samples contained a
small percentage of pets (i.e., cats and dogs; 3.3% ± 0.6%; Table 1).
Percentages of rodents and pets in fecal samples were higher in
high-density sites and percentages of deer were higher in low-
density sites (Fig. 3a). Mammals were more common in the diet in
December, March, April, and June than in August or October and
more common in the diet in high-density sites than low-density
sites throughout the year, especially in August, October, and
March (Fig. 3b). Pet hairs were least common in the diet in December
(Fig. 4), the time of year when coyote–pet conflicts were highest in
the DMA (see Fig. 3bin Poessel et al. 2013). Month was not a
significant variable in the ANOVA testing for a relationship be-
Fig. 3. Mean (±SE) percentage of hairs of mammal groups found in coyote (Canis latrans) fecal samples in high- and low-density sites (a) and
mean (±SE) mass of all hairs found in coyote fecal samples by month in high- and low-density sites (b) in the Denver metropolitan area, from
2013 to 2014. The “unknown” category in panel aincludes all unidentified mammal hairs found in coyote fecal samples during the study.
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tween the amount of pet hair in the diet and month (F
= 1.8,
P= 0.123).
Fecal samples contained 33 different species of seeds that coy-
otes either fed on directly or were present in prey items of coyotes.
Russian olive (Elaeagnus angustifolia L.), cactus (species of the genus
Opuntia Mill.), corn (Zea mays L.), field bindweed (Convolvulus arvensis L.),
sorghum (species of the genus Sorghum Moench), American plum
(Prunus americana Marshall), and sunflower (Helianthus annuus L.)
were most commonly present in fecal samples by frequency
(Table 1). Russian olive and grapevine (species of the genus Vitis L.)
were more common in high-density sites, whereas cactus, choke-
cherry (Prunus virginiana L.), and corn were more prevalent in low-
density sites by mass (Fig. 5a). The presence of seeds in fecal
samples also reflected seasonal availability, with more plants con-
sumed in August and October than in December, March, April, or
June. Fecal samples also contained a higher amount of seeds in
August and October in low-density sites than in high-density sites
(Fig. 5b), especially of native plants.
Ground beetles and grasshoppers made up the majority of exo-
skeletons found in scats. The presence of exoskeletons was high-
est in August and October and lowest in March and April (Fig. 6a).
Exoskeletons were also more common in low-density sites than
in high-density sites for most of the year (Fig. 6a). The amounts of
feathers in scats varied across months in both high- and low-
density sites and were most common in December and April
(Fig. 6b). Feathers were also more common in high-density sites
than in low-density sites for most of the year (Fig. 6b).
Coyotes appeared to have consumed anthropogenic food items
in low amounts in our study area. Pets comprised only 3% of
mammal hairs in the coyote diet, and they were more commonly
found in high-density sites (Fig. 3a). This finding corroborates
other studies that have determined that coyotes consume domes-
tic cats and dogs in low quantities (Fedriani et al. 2001;Morey et al.
2007;Gehrt and Riley 2010;Lukasik and Alexander 2012). Murray
et al. (2015) also found that coyotes in higher density (urban) areas
consumed more pets than did coyotes in lower density (rural)
areas. In Tucson, Arizona, Grubbs and Krausman (2009) observed
19 cats that were killed by coyotes, 18 of which were consumed.
However, they did not report a dietary analysis for these coyotes,
so the importance of cats in the coyote diet in this study area is
In the DMA, over 92% of coyote–human conflicts reported be-
tween 2003 and 2010 were incidents with pets (471; Poessel et al.
2013). These included injuries and deaths of cats and dogs caused
by coyotes, although we did not have information on how many of
these pets were killed. Hence, coyotes can be a serious threat to
the safety of domestic pets. Furthermore, coyote conflicts with
pets were more frequent during the winter months (December
and January; see Fig. 3bin Poessel et al. 2013). However, pets com-
prised only a very small percentage of the coyote diet in our study,
and pet hair did not increase in the diet during winter, possibly
indicating that coyotes usually do not consume the cats and dogs
that they kill. These results support the hypothesis that coyote
conflict with pets is primarily driven by competition or a threat
response, rather than predation. Coyotes may view cats as com-
petitors because cats will hunt rodents, a primary component of
the coyote diet, similar to the finding that coyotes also perceive
other canids, such as dogs and foxes, as competitors for food and
habitat (Gosselink et al. 2003). Winter months coincide with coy-
ote breeding season, so during this time of year, coyotes may view
dogs as potential competitors for mates, leading to increased con-
flicts with pets (Poessel et al. 2013). Additionally, pets increased in
the coyote diet in March, the time of year when births of pups
Fig. 4. Mean (±SE) percentage of hairs from pets found in coyote (Canis latrans) fecal samples in high- and low-density sites by month in the
Denver metropolitan area, from 2013 to 2014.
Poessel et al. 293
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begin, so coyotes may perceive dogs as a threat to their pups and
pets killed during this time also may be consumed.
We found only a small amount of trash in the coyote diet. Only
0.3% of fecal sample masses consisted of trash in both high- and
low-density sites. However, 27 of 64 fecal samples (42%) contained
trash items (Table 1), although we do not know how many individual
scats contained trash. Hence, although coyotes do not appear to be
consuming large amounts of garbage, they may be accessing it fre-
quently, even though during our study we did not observe coyotes
eating curbside trash or entering trash dumpsters.
We used trash to measure anthropogenic food in the coyote diet
because this human-associated item was indigestible and easily
recovered in fecal samples. However, coyotes could have con-
sumed other anthropogenic food items, such as pet food or hu-
man food that is digested and cannot be measured in scats. Thus,
fecal analysis has limitations for examining the diet of wildlife
species in urban environments. Stable isotope analysis is an alter-
native method that has been used in urban systems to examine
coyote diets (Murray et al. 2015;Newsome et al. 2015). This tech-
nique is better able to discriminate between natural prey items
and anthropogenic resources by estimating the isotopic composi-
tion of each food item and of coyotes, measured in hair or vibris-
sae. Previous studies using this method have shown that some
coyotes in urban areas will consume anthropogenic food (Murray
et al. 2015;Newsome et al. 2015), and stable isotope analysis can
report a higher prevalence of anthropogenic foods in coyote diets
than scat analysis. Therefore, if assessing anthropogenic food re-
sources in the urban coyote diet is a primary objective, then fu-
ture studies should include this technique or others like it (e.g.,
fatty acid analyses) in conjunction with fecal analysis to obtain a
more accurate assessment of coyote diets in urban areas.
The diet of coyotes in our study area varied both spatially and
temporally, and coyotes showed evidence of opportunistic forag-
ing. Generally, coyote diets consisted of more mammals in high-
density sites and more plants in low-density sites, although some
individual animal and plant species varied from this pattern. Like-
Fig. 5. Mean (±SE) mass of seeds found in coyote (Canis latrans) fecal samples in high- and low-density sites of the most common plant
species (a) and all plants by month (b) in the Denver metropolitan area, from 2013 to 2014. The “other” category in panel aincludes all other
plant species consumed by coyotes during the study.
294 Can. J. Zool. Vol. 95, 2017
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wise, mammals and birds were more common in winter and
spring (December–June), whereas plants and invertebrates were
more prevalent in late summer and autumn (August–October),
likely due to seasonal food availability. Coyotes in our study area
preferred natural habitat over developed landscapes, although
they did use residential areas, especially at night (Poessel et al.
2016); however, we do not know where coyotes were primarily
Specifically for animals, mammal hair, particularly that of ro-
dents and rabbits, was the most prevalent diet item in coyote fecal
samples. Coyotes consumed rodents more often in high-density
sites and deer more often in low-density sites (Fig. 3a). Mule deer
(Odocoileus hemionus (Rafinesque, 1817)) require large amounts of
space, are dependent on natural areas, and therefore were more
readily available in the low-density sites than in the high-density
sites. Thus, coyotes may have increased rodents in their diet in
high-density areas because deer were not as available.
Coyote consumption of rodents followed the seasonal pattern
of higher prevalence in winter and spring, although rodents were
common in the diet throughout the year. Because every one of the 64
fecal samples contained rodent hair, this mammal group influ-
enced the overall seasonal pattern for all mammals. Deer hair was
most common in fecal samples in June, after the birthing period
of fawns when they are vulnerable to coyote predation. However,
coyotes fed on deer throughout the year, likely available as car-
rion from roadkill. Lagomorphs were more prevalent in the diet in
late autumn and early winter than in spring and summer. Morey
et al. (2007) also found that lagomorphs occurred in coyote diets in
lower frequencies during the summer, suggesting that rabbits are
better able to avoid capture by coyotes during a time of increased
vegetation growth. Furthermore, fruit is more available during
summer, so coyotes may be switching from animals to plants
during this time.
Specifically for plants, coyotes consumed Russian olive (non-
native) and grapevine (non-native) more often in high-density
sites and cactus (native), chokecherry (native), and corn (non-
native) more often in low-density sites (Fig. 5a). Russian olive, a
highly invasive species, has become naturalized throughout the
western United States and is primarily found in riparian areas
(Shafroth et al. 1995). The high-density sites in our study area were
located near riparian areas, and coyotes used riparian areas fre-
quently (Poessel et al. 2016), leading to a high prevalence of this
Fig. 6. Weighted average scores of exoskeletons (a) and feathers (b) found in coyote (Canis latrans) fecal samples by month in high- and
low-density sites in the Denver metropolitan area, from 2013 to 2014.
Poessel et al. 295
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plant in coyote scats at these sites. Grapevines are widely culti-
vated in gardens, likely resulting in a large number of these plants
in residential areas and a high occurrence of these seeds in coyote
scats at high-density sites. Cactus and chokecherry are both native
species to Colorado and were common in low-density sites where
coyotes were more likely to consume native fruits. Finally, corn
may have been more common in low-density sites because of the
higher presence of deer in these areas; we speculate that residents
in these sites may be placing corn outside to feed deer, which can be
accessed by coyotes. Corn fields also are present near the low-
density sites, and we occasionally observed coyotes in these fields.
Coyote consumption of most plant species followed the same
seasonal pattern of high prevalence in summer or autumn. Russian
olive was found in the coyote diet in higher masses in August,
October, and December. However, corn was most commonly
found in the diet in early winter (December), a time of year when
coyotes might be more food-stressed (Bekoff and Wells 1981).
Corn, as well as sorghum, is found in different brands of birdseed,
and residents of the DMA may be leaving out dried or cracked corn
and birdseed in bird feeders. Thus, coyotes likely were utilizing
this food source to supplement their diet in the winter. Although
we can only speculate, the presence of corn in the coyote diet is
likely anthropogenically driven, whether from residents feeding
deer, placing corn in bird feeders, or cultivating corn plants.
Russian olive was the most frequently consumed plant, occur-
ring in 17 of 64 fecal samples (27%; Table 1). Although coyotes
consumed 14 different non-native plant species, the largest vol-
ume of seeds in the coyote diet consisted of native species (82% of
total seed mass, although the majority of this mass was from only
two species; Table 1). Hence, coyotes may be an important dis-
perser of not only Russian olive, but also a variety of native plant
species in this highly urbanized area.
Our scat analysis revealed that pets consisted of only a small
percentage of the coyote diet, and pet consumption decreased
when coyote–pet conflicts increased, which supports the hypoth-
esis that coyote conflict with pets is primarily driven by competi-
tion or a threat response, rather than predation. Coyotes in urban
areas have a wide variety of foods available to them, so they do not
need to consume pets. However, territoriality in coyotes is strong
and they will remove perceived threats to their ability to survive
and reproduce. Hence, pet owners should be diligent in their
efforts to protect their pets by not letting pets outside unsuper-
vised, including in their yards, by keeping pets on leash when
walking them in natural areas, and by being especially aware of
coyotes during winter. Our work further revealed that trash did
not constitute a significant portion of the diet of coyotes, but our
analysis technique was not sensitive to detecting certain anthro-
pogenic food sources that are difficult to detect with scat analysis.
Urban residents can reduce the amount of such food available to
coyotes by securing trash and removing pet food from outdoor
areas. As reports of wildlife conflicts in urban areas continue to
increase, understanding how urban-adapted species, such as coy-
otes, use anthropogenic resources may provide insight that can be
used to promote coexistence between humans and wildlife.
We thank the many technicians and volunteers who assisted
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S. Smith, R. Galvan, L. Bowie, M. Casey, J. Hawkins, C. Mitchell,
B. Tomlin, P. Mills, R. Deangelis, and C. Kerr. Funding and logisti-
cal support were provided by the U.S. Department of Agriculture,
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... For example, Fedriani et al. (2001) also found high dietary diversity in an urban study location in the Santa Monica Mountains, and Larson et al. (2015) found mammalian remains in 80% of coyote scats in San Diego, California. Our findings were also generally consistent with studies outside of California, where rabbits were also the primary prey of coyotes in Madison, Wisconsin (Garwood et al. 2015), and the second highest prey in Denver, Colorado after rodents (Poessel et al. 2017). While researchers in Calgary, Alberta, Canada found urban coyotes ate a predominately natural diet with a similar 14% occurrence of anthropogenic food items, they only found domestic pets in less than 2% of scats (Lukasik and Alexander 2012). ...
... This may indicate that relatively small natural habitat patches within heavily developed urban areas allow for the maintenance of natural behaviors of opportunistic hunting and foraging. Some urban area studies have also found similar seasonal variations in mammalian and vegetation consumption, including in the Denver area (Poessel et al. 2017) and Chicago (Morey et al. 2007). In Los Angeles, Larson et al. (2020) found native fruits in less than 2% across sites and seasons, but their ornamental fruit results were higher, with an FO of up to 32% at their suburban site in the dry season. ...
... Intraguild predation is common in carnivores, and coyotes are among the top three canids to practice this behavior (Palomares and Caro 1999), killing bobcats Lynx rufus, and grey foxes Urocyon cinereoargenteus (Fedriani et al. 2000), as well as domestic and feral canids and felids (Gehrt et al. 2013, Kays et al. 2015, Poessel et al. 2017. After domestic cats, raccoons were the second most common mesocarnivore found in scats in this study but had low occurrence at 6% compared to 18% found in rural Ohio (Cepek 2004) or 27% in urban New York (Henger 2022). ...
Full-text available
Coyotes Canis latrans in urban landscapes provide important food web functions and ecological services but can also trigger human-wildlife conflict when their diet includes anthropogenic resources or domestic pets. As adaptable omnivores, coyotes adjust their diet to their environment, routinely switching among food items to accommodate spatial and seasonal differences in availability. To evaluate the coyote’s potential impacts within the food web of urban Long Beach, California where human-wildlife conflict involving coyotes may occur, we analyzed 115 scat samples collected once every two weeks from four open space fragments inside the urban matrix. We hypothesized that differences in scat composition would correlate with seasonal and site differences, with greater use of anthropogenic resources during the dry season supplementing lower prey availability, and with greater consumption of wild mammal prey during the wet season when fruiting plants are less abundant. We found coyote diet was predominately composed of natural prey and vegetation year-round, with seasonal variation. Mammals made up more of coyote diet in the wet season than the dry, while invertebrates and vegetation were more prevalent in dry season scats. Coyotes relied on rabbits as their main prey year-round across all sites. Domestic cats Felis catus were the third most common individual prey species found in coyote scats, occurring in 14% of scat samples in both seasons. Coyotes also supplemented seasonally available natural food sources with anthropogenic resources, which occurred in 13% of coyote scats overall with no significant seasonal variation. While rabbits appeared in scat from all sites, the occurrence of invertebrates, small mammals, and vegetation in scats varied between sites. While there is a potential for human-wildlife conflict in coyote’s consumption of feral or domestic cats, coyotes may also be providing an ecological service by reducing cats in natural habitat fragments.
... We determined scats to be coyote based on the location, color, size, shape, and amount of hair present (Cypher F I G U R E 2 Coyote scat (A) and complete Russian olive seeds extracted from scat (B) found in western North Dakota, USA, 2020. 1991, Poessel et al. 2017). Scats collected contained bone fragments and hair consistent with coyote scats. ...
... Scats collected contained bone fragments and hair consistent with coyote scats. No human dwellings are located near any of our sites and domestic dog scats generally lack hair, bones, and seeds (Poessel et al. 2017). At site 1, we collected North American porcupine (Erethizon dorsatum) scats, which also contained Russian olive seeds. ...
Russian olive (Elaeagnus angustifolia) is an invasive tree that has spread throughout much of the western United States. The mode of seed dispersal occurs by hydrochory and possibly by birds. Seed dispersal by frugivorous mammals has not been investigated. Between 15 October and 4 November 2020, we walked through Russian olive windbreaks in western North Dakota, USA, and surveyed for mammal scat, and found 10 coyote (Canis latrans) and 54 porcupine (Erethizon dorsatum) scats that contained intact Russian olive seeds. We subsequently evaluated the viability, germination frequency, and time to germination of seeds ingested by coyote and porcupine relative to un‐ingested control seeds harvested from trees at sites where we collected scat. Overall, Russian olive seeds that passed through mammal intestinal tracts had similar viability and equivalent (porcupine) or higher (coyote) germination frequency compared to controls. Additionally, coyote‐ingested seeds germinated earlier (time to germination was low) than controls, but porcupine‐ingested seeds were similar to controls. Thus, our data supports the idea that mammals may be agents of regional Russian olive seed dispersal. Russian olive seeds can be dispersed via endozoochory by mammals. Thus, mammals could play a role in spreading Russian olive into areas where other modes of transport (e.g., hydrochory) cannot.
... Coyotes (Canis latrans) are one example of a highly successful urban species that has received considerable attention from urban wildlife managers. The foremost adaptations that have enabled coyotes to succeed in urban areas include broadening their diet to include more anthropogenic food (Gehrt and Riley 2010;Murray et al. 2015a;Poessel, Mock, and Breck 2017) and shifting to a more nocturnal lifestyle to avoid human activity (Grinder and Krausman 2001;Gese, Morey, and Gehrt 2012;Murray and St. Clair 2015). Anthropogenic food supplementation has created numerous challenges for urban coyote management, as food supplementation has been connected to poor health (Sugden et al. 2020), human-wildlife conflict (Murray et al. 2015a) and a higher prevalence of parasites (Murray et al. 2016). ...
... For example, anthropogenic food and domestic pets have been estimated to comprise up to 80% of the diet of urban coyotes in southern California (Larson et al. 2020), but this proportion is only 40-50% in Edmonton (Murray et al. 2015b) and 20-35% in Chicago (Newsome et al. 2015). This broad range from isotopic data parallels the 40-80% prevalence of anthropogenic food reported in scat-based analyses (Murray et al. 2015a;Poessel, Mock, and Breck 2017;Larson et al. 2020). Further variation may stem from disease status: diseased coyotes consumed more anthropogenic food and less prey than healthy coyotes (Murray et al. 2015b). ...
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Many generalist species thrive in urban environments by supplementing their diets with anthropogenic food, which creates numerous challenges for managing urban wildlife. Management could be advanced with more information on how spatial and temporal variation in habitat use by urban animals predicts variation in their dietary ecology. In this study, we used stable isotope analysis complemented with GPS collar location data to determine how diet composition and the dietary niche of coyotes (Canis latrans) varied across a sample of 169 individuals collected along an urban-to-rural gradient in Alberta, Canada. We further categorized urban individuals as either matrix (frequent use of developed areas) or greenspace (use of natural areas) via GPS locations. Matrix coyotes were isotopically distinct from all other coyote populations: they had the largest dietary niche, exhibited the most among-individual variation in diet, consumed the most anthropogenic food and fruit, and consumed the least amount of prey. Greenspace coyotes consumed more anthropogenic food than rural and suburban coyotes but otherwise exhibited similar niche width, among-individual heterogeneity, and prey consumption. We further tested for seasonal dietary variation and found that urban coyotes had a larger dietary niche during the summer, when they also consumed more anthropogenic food. Our conclusions were robust to our choice of mixing model parameters, including discrimination factors, suggesting that these methodological choices have limited effect when discerning relative trends among populations. Overall, our results suggest that management of urban coyotes should target the food sources accessible to coyotes in the urban matrix to reduce human–coyote conflict.
... Other studies carried out in Durango reported similar food preferences, adding insects as an important food item during the dry season (Servín and Huxley 1991;Grajales-Tam and González-Romero 2014). Diet preferences coincide with the ecosystem role this canid plays as a biological control of rodent and lagomorphs populations, and seed disperser in different ecosystems in which it is distributed (Servín and Huxley 1991;Grajales-Tam et al. 2003;Cruz-Espinoza et al. 2008;Arias-Del Razo et al. 2011;Grajales-Tam and González-Romero 2014;Poessel et al. 2017). ...
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The coyote (Canis latrans) is a widespread predator with a high degree of adaptation to different ecosystems. The objective of this study was to estimate the relative abundance index, habitat selection, and diet of C. latrans through scent stations, in two types of vegetation located in northwestern Chihuahua. From April 2018 to March 2019, ten fixed scent stations (SS) were placed in pine-oak forest and other ten in semi-open grassland, with a linear separation of 500 meters between each station to achieve a five km transect in each vegetation during 12 samplings (two sampling night per month) for totaling 420 SS after discarding inactive SS. The relative abundance index of coyote showed that both types of vegetation, pine-oak forest (0.30) and semi-open grassland (0.23) were used in a similar way. The habitat selection test (Chi2) showed that coyote abundance and type of vegetation were independent (χ2 = 2.96, P > 0.05), not showing statistically significant differences in annual relative abundance index of coyotes between the two vegetation types. The food items detected in thirty-four scats collected belonged to mammals (55.9 %), fruits (35.3 %) and arthropods (8.8 %). Rodents and lagomorphs were the main source of food. Throughout the sampling period, the pine-oak forest showed the greatest relative abundance index of C. latrans. This may be because the forest provides them with shelter from climatic situations and a greater variety of seeds, fruits and insects. In this study the two seasons with the highest relative abundance were spring and autumn in both ecosystems, coinciding with an increase in rainfall, resulting in an abundance of potential prey (rodents and lagomorphs), and other food items like fruits. Regarding annual diet the most consumed prey were mammals. We found differences in diet between seasons, that can be explained by the variation in food availability among seasons.
... Urbanization is an important contributor to the rise in HWC worldwide and the growth of human-carnivore conflicts in North America. Certain species such as coyotes demonstrate generalist tendencies in their foraging (e.g., use of both anthropogenic and natural food sources) and habitat requirements (e.g., use of natural cover patches within residential neighborhoods), making them well-suited to adapt to urban environments (Poessel, Breck, & Gese, 2016;Poessel, Mock, & Breck, 2017). Coyotes may also be more predisposed than some other carnivores to survive in, and even newly colonize, urban areas due to their heightened ability to tolerate human presence (Sol, Lapiedra, & Gonzalez-Lagos, 2013). ...
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Human-wildlife conflict ( HWC ) is a global phenomenon with serious implications for biodiversity conservation and human well-being. Innovative solutions demand greater attention to the social factors contributing to HWC , including human thought and behavior, which can be examined through the lens of social psychology. Using the case of human-coyote conflict in North America, this study employed a mixed-methods social psychology approach to explore the potential for citizen science to serve as a tool for HWC mitigation. Quantitative surveys and interviews with volunteers in a coyote-focused citizen science program in Colorado revealed that the program is positively affecting participants’ attitudes/beliefs and empowering them to take action to address conflicts in their communities. The article concludes with recommendations for future evaluative research in this area as well as ways to more generally improve social-psychology applications in wildlife conservation.
... Despite their widespread distribution, information on Coyote population parameters (e.g., density, abundance), life history traits (e.g., survival, recruitment), diet, and habitat use in both urban and rural environments is scarce for many areas (Poessel et al. 2017, Scotten 2019. ...
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Coyotes (Canis latrans) are expanding their range and due to conflicts with the public and concerns of Coyotes affecting natural resources such as game or sensitive species, there is interest and often a demand to monitor Coyote populations. A challenge to monitoring is that traditional invasive methods involving live-capture of individual animals are costly and can be controversial. Natural resource management agencies can benefit from contemporary noninvasive genetic sampling approaches aimed at determining key aspects of Coyote ecology (e.g., population density and food habits). However, the efficacy of such approaches under different environmental conditions is poorly understood. Our objectives were to 1) examine accumulation and nuclear DNA degradation rates of Coyote scats in metropolitan and rural sites in Florida to help optimize methods to estimate population density; and 2) explore new genetic methods for determining diet of Coyotes based on vertebrate, plant, and invertebrate species DNA identified in scat. Recently developed DNA metabarcoding approaches make it possible to simultaneously identify DNA from multiple prey species in predator scat samples, but an exploration of this tool for assessing Coyote diet has not been pursued. We observed that scat accumulation rates (0.02 scats/km/day) did not vary between sites and fecal DNA amplification success decreased and genotyping errors increased over time with exposure to sun and precipitation. DNA sampling allowed us to generate a Coyote density estimate for the urban environment of eight Coyotes per 100 km2, but lack of recaptures in the rural area precluded density estimation. DNA metabarcoding showed promise for assessing diet contributions of vertebrate species to Coyote diet. Feral Swine (Sus scrofa) were detected as prey at higher frequencies than previously reported. We identify several considerations that can be used to optimize future noninvasive sampling efforts for Coyotes in the southeastern United States. We also discuss strengths and drawbacks of utilizing DNA metabarcoding for assessing diet of generalist carnivores such as Coyotes. .
... These species have become particularly adapted to urban environments; red fox can be found in major cities worldwide, and coyotes have been seen in every large city in the continental United States (21,22). Studies of urban adapted coyotes have found that they are less likely to flee from humans, more likely to engage in conflicts with pets, and more likely to prey on pet dogs and cats than coyotes in rural environments, all behaviors that could increase the coyotes' risk of being exposed to SARS-CoV-2 (22,23). ...
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Severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) has proven to be a promiscuous virus, capable of infecting a variety of different animal species, but much work remains in determining the susceptibility of common wildlife species to the virus. Here, we demonstrate that following experimental inoculation with SARS-CoV-2, red fox ( Vulpes vulpes ) become infected and can shed virus in oral and respiratory secretions. Conversely, experimentally challenged coyotes ( Canis latrans ) did not become infected or shed virus. Our results add red fox to the animal species known to be susceptible to SARS-CoV-2 and suggest that they may contribute to continued maintenance and transmission of the virus. Article Summary Line Experimental infection of red fox ( Vulpes vulpes ) and coyotes ( Canis latrans ) with SARS-CoV-2 revealed that red fox are susceptible to infection and can shed virus, while coyotes do not become infected.
... The variation in the frequency of consumption in the trophic categories that make up the diets of these two canids shows their ability to adapt to different habitat conditions. In the case of the coyote, it has been reported that this species responds to changes in resource availability by modifying their preferences when an important food source becomes less abundant [20,34,108,109]. This also appears to occur with gray foxes, since they have also demonstrated their adaptability to changes in the availability of food resources, whether due to stochastic events, temporal variation, or differences in the habitat types they occupy [18,20,23,24,102]. ...
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Resource partitioning, and especially dietary partitioning, is a mechanism that has been studied for several canid species as a means to understand competitive relationships and the ability of these species to coexist. Coyotes ( Canis latrans ) and gray foxes ( Urocyon cinereoargenteus ) are two canid species that are widely distributed, in Mexico, and they are sympatric throughout most of their distribution range. However, trophic dynamic and overlap between them have not been thoroughly studied. In order to better understand their ecological relationship and potential competitive interactions, we studied the trophic niche overlap between both canids in a temperate forest of Durango, Mexico. The results are based on the analysis of 540 coyote and 307 gray fox feces collected in 2018. Both species consumed a similar range of food items, but the coyote consumed large species while the gray fox did not. For both species, the most frequently consumed food categories throughout the year and seasonally were fruit and wild mammals (mainly rodents and lagomorphs). Coyotes had higher trophic diversity in their annual diet ( H’ = 2.33) than gray foxes ( H’ = 1.80). When analyzing diets by season, trophic diversity of both species was higher in winter and spring and tended to decrease in summer and autumn. When comparing between species, this parameter differed significantly during all seasons except for summer. Trophic overlap throughout the year was high ( R 0 = 0.934), with seasonal variation between R 0 = 0.821 (autumn) and R 0 = 0.945 (spring). Both species based their diet on the most available food items throughout each season of the year, having high dietary overlap which likely can lead to intense exploitative competition processes. However, differences in trophic diversity caused by differential prey use can mitigate competitive interactions, allowing these different sized canid species to coexist in the study area.
... However, urban ecosystems are more open than most natural systems, importing most food resources from hundreds or thousands of miles away. Although urban consumers may continue to draw on resources from natural sources, consumers in natural compartments may become more reliant on novel food sources if the natural environment degrades (Poessel et al. 2017). ...
Ecosystems are defined, studied, and managed according to boundaries constructed to conceptualize patterns of interest at a certain scale and scope. The distinction between ecosystems becomes obscured when resources from multiple origins cross porous boundaries and are assimilated into food webs through repeated trophic transfers. Ecosystem compartments can define bounded localities in a heterogeneous landscape that simultaneously retain and exchange energy in the form of organic matter. Here we developed and tested a framework to quantify reciprocal reliance on cross‐boundary resource exchange and calculate the contribution of primary production from adjacent ecosystem compartments cycling through food webs to support consumers at different trophic levels. Under this framework, an integrated ecosystem can be measured and designated when the boundary between spatially distinct compartments is permeable and the bidirectional exchange of resources contributes significantly to sustaining both food webs. Using a desert river and riparian zone as a case study, we demonstrate that resources exchanged across the aquatic‐riparian boundary cycle through multiple trophic levels. Further, predators on both sides of the boundary were supported by externally produced resources to a similar extent, indicating this is a tightly integrated river‐riparian ecosystem and that changes to either compartment will substantially impact the other. Using published data on lake ecosystems, we demonstrated that benthic and pelagic ecosystem compartments are likely not fully integrated, but differences between lakes could be used to test ecological hypotheses. Finally, we discuss how the integrated ecosystem framework could be applied in urban‐preserve and field‐forest ecosystems to address a broad range of ecological concepts. Because few systems function in complete isolation, this novel approach has application to research and management strategies globally as ecosystems continue to face novel pressures that precipitate cascading ecological repercussions well beyond a bounded system of focus.
... Lukasik and Alexander (2011) found that coyote attacks on pets were more frequent in neighborhoods where coyotes had a greater proportion of anthropogenic food in their diets. However, Poessel et al. (2017b) found that seasonal increases in the frequency of coyote attacks on pets did not result in a seasonal increase in the presence of pets in a coyote's diet. This suggests that coyotes attack pets because they are either perceived as competitors for food resources or as a threat to their young, not because the pets themselves are perceived as a prey item. ...
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Coyotes (Canis latrans) have established populations in most major urban centers across North America. While the risk of attacks on humans or their pets is low, the presence of carnivores in areas with high human use has resulted in increased public concern. Having a clearer understanding of which animals are more likely to interact with humans and when interactions are more likely to occur will help mitigate human-carnivore conflicts. Despite clear broad-scale patterns of human avoidance, human-coyote interactions occur most frequently in residential areas. Our purpose was to determine if use of residential areas varied consistently across individuals or time. We used locations from GPS collars deployed on 14 coyotes in the Greater Toronto Area, Ontario, Canada from 2012 to 2017 to fit a step selection function. Average (±SE) home range size estimates were 17.3 ± 4.6 km 2 for resident coyotes and 102.8 ± 32.9 km 2 for non-residents. We found that coyotes used natural areas more (β = 0.07, SE = 0.02, p < 0.0001), and roads (β = −0.50, SE = 0.13, p < 0.0001) and residential areas (β = −0.79, SE = 0.21, p = 0.0001) less during the day than at night. We also found that coyotes were more likely to use residential areas in the breeding season from January to April (β = 0.69, SE = 0.20, p = 0.0007) and the pup rearing season from May to August (β = 0.54, SE = 0.13, p < 0.0001) than in the dispersal season from September to December. Lastly, we found that resident coyotes were less likely to use residential areas than non-resident coyotes (β = −1.13, SE = 0.26, p < 0.0001). As far as we are aware, our study is the first to identify the seasons when coyotes are more likely to use residential areas. The seasonal patterns in habitat use that we observed reflect patterns that have been previously reported for human-coyote conflicts. Our results demonstrate that reducing the availability of anthropogenic food sources in residential areas, particularly in the winter and spring, should be a priority for managers aiming to reduce human-coyote conflict in urban areas.
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The increase of global urbanization can have effects on wildlife species, including carnivores such as coyotes (Canis latrans). As coyotes continue to settle in more urban areas, reports of human-coyote conflicts , such as attacks on humans or pets, may also increase. Understanding environmental variables that might influence whether or not coyotes and human-coyote conflicts will occur in certain urban areas may assist wildlife officials in creating management plans for urban wildlife. We conducted a survey of 105 urban areas in the United States requesting information on the occurrence of coyotes and human-coyote conflicts. We analyzed the responses with data on human population size, geographic region, land cover, housing density, and precipitation. Larger urban areas were more likely to contain both coyotes and human-coyote conflicts, and were also more likely to have greater numbers of conflicts. Urban areas in the western regions with larger amounts of high-intensity development and less forested and agricultural areas were more likely to have conflicts. Most urban areas considered the management of conflicts to be of low priority and emphasized education of citizens rather than removal of individual coyotes. Our results may assist urban wildlife managers in understanding the geographic and demographic factors correlated with the occurrence of coyotes and human-coyote conflicts. Practices such as education campaigns and landscape design incorporating wildlife habitat modifications (e.g., reducing dense cover) may reduce human-carnivore conflicts in urban ecosystems.
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Urbanization alters landscapes and ecosystem processes that result in negative impacts for many species. However, urbanization also creates novel environments that certain species, including carnivores, are able to exploit. Coyotes (Canis latrans) are 1 example of a species capable of exploiting urban environments throughout North America and, in some cases, becoming involved in human–coyote conflict. As part of a comprehensive study of human–coyote coexistence in the Denver metropolitan area of Colorado, we investigated the spatial ecology of coyotes to determine movement and activity patterns relative to the urban matrix. We examined home-range size, habitat use, and resource selection for 22 coyotes monitored with GPS collars during 2012–2014. Mean (± SD) home-range size of resident coyotes (11.6±11.0 km2) was smaller than ranges of transient coyotes (200.7±232.4 km2). Home-range size did not vary by season or sex, but resident coyotes during the day (7.2±10.5 km2) had smaller home ranges than during the night (11.3±10.8 km2). Coyotes had high percentages of developed lands (44.5±18.9%) within their home ranges, contrary to previous studies of urban coyotes. However, the percentage of coyote locations in natural lands (48.9±22.4%) was higher than in developed lands (20.6±11.7%). Home-range size of residents was not related to either the percentage of developed lands or altered lands within home ranges. Coyotes selected natural lands over developed lands, and they increased activity at night. Although coyotes were able to thrive in home ranges containing large amounts of development, they continued to avoid areas with high human activity by primarily residing in areas with natural land cover. Similar to urban areas throughout the Northern Hemisphere, coyotes in the Denver metropolitan area have become efficiently adapted to a highly developed landscape, reflecting the flexible nature of this opportunistic carnivore.
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To evaluate whether the abundance of coyotes Canis latrans was influenced by the availability of anthropogenic foods in a humanized landscape, we compared three neighboring areas (hereafter referred to as NA, CA, and SA) under contrasting human pressures within the Santa Monica Mountains of California, USA. We quantified the use of anthropogenic foods by coyotes and assessed local densities within these three regions. Overall, 761 coyote feces were analyzed; identified food items were categorized into 11 food types (7 native and 4 anthropogenic). Though small mammals (lagomorphs and rodents) were the main prey of coyotes in all areas and seasons, log-linear modeling of multiway contingency tables indicates that consumption of anthropogenic foods by coyotes varied significantly throughout study areas. Thus, in the most humanized area (CA; 24% of this region is residential habitat), anthropogenic foods (trash, livestock, domestic fruit) comprised seasonally between 14 and 25% of total items in coyote diets, whereas in the least humanized area (NA; 2% residential) anthropogenic foods only comprised seasonally between 0 and 3% of items. Coyote density, estimated by foot-hold trapping surveys and by genotyping feces, was also highly variable between areas. The heavily human-impacted CA area had the highest coyote density (2.4-3.0 ind. km-2), whereas coyote density was significantly lower (0.3-0.4 ind. km-2) in the least humanized area (NA). In the third region (SA; 10% residential), with an intermediate level of human pressure, both importance of anthropogenic foods in coyote diet (4-6%) and coyote density (1.6-2.0 ind. km-2) were intermediate compared to the other regions. Our data suggest that subsidization by anthropogenic foods augments coyote densities and alters their diets in the Santa Monica Mountains, California. We include data from literature to show that anthropogenic foods are used by omnivorous mammals throughout the world. Surprisingly, however, the potential effects of allochthonous inputs on such species are not well-understood. Thus, further research on this phenomenon in humanized landscapes is needed.
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Reports of encounters between people and generalist urban-adapted carnivores are increasing around the world. In North America, coyotes Canis latrans are among the carnivores that appear to be especially capable of incorporating novel anthropogenic food types, including those found in cities. Consuming anthropogenic food may benefit coyotes by increasing their dietary diversity, but it may also lead to increased interactions and conflicts with humans. To test these hypotheses, we compared the diets of urban and rural coyotes from two urban and three rural sites spanning 32 200 km2 in Alberta, Canada. We analyzed scat samples to calculate diet diversity at the level of both individuals (species per scat) and populations (Shannon index) and to determine the frequency of anthropogenic food consumption. We complemented this comparison with stable isotope analyses of hair samples taken from individual urban and rural coyotes that were or were not reported by the public for repeatedly visiting backyards and schoolyards during the day. Relative to rural coyotes, urban coyotes had more diverse diets at the level of both individuals and populations, consumed anthropogenic food more often, and animals less often, than rural coyotes. Although urban coyotes assimilated more anthropogenic food than the rural coyotes overall, the urban coyotes reported for conflict assimilated less protein and were more likely to be diseased. Our results suggest that processed anthropogenic food may contribute to the success of urban coyotes, but does not entirely correlate with conflict. Instead, some seemingly innocuous, but low-protein food sources such as bird feeders, compost, and cultivated fruit trees may contribute disproportionately to encounters with people for coyotes and other urban-adapted opportunistic carnivores.
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Eighteen radio-equipped coyotes (Canis latrans) were found to have home ranges averaging 68.7 km2 for adult females, 31.3 km2 for adult males, 39.9 km2 for yearling females, and 1.0 km2 for pups. Coyotes involved in pup rearing occupied smaller home ranges than did unmated coyotes. Home ranges of adult females overlapped, as did adjacent male-female home ranges. Coyotes were located more in savanna and less in prairie than expected (P < 0.05) by the availability of these habitats. Seventy-eight percent of the observations of coyotes were of lone animals, 17% were of pairs, and less than 6% were of more than 2 coyotes. Analysis of coyote scats indicated rodents, fruits and seeds, and fawn deer (Odocoileus virginianus) were important foods.
The coyote (Canis latrans) is a common resident in urban areas throughout the United States, yet little is known about coyote diets in these environments. I characterized the annual diet of coyotes in an urban environment of western Washington by analyzing their scat from three areas representing typical patterns of human occupation and density: residential (1413 humans/km2), mixed agricultural residential (348 humans/km2), and mixed forest-residential (126 humans/km2). Coyote scats were collected twice a month for 1 year (Nov. 1989-Oct. 1990) in each habitat type. Fruits and mammals were the largest classes of food items in all habitat types and their seasonal use was similar among habitats. Apple (Malus spp.) and cherry (Prunus spp.) were the most abundant fruits in the scats, and ranged from 22-41% and 9-13% of the annual diet, respectively. Vole (Microtus spp.) was the most abundant mammalian food item (41.7%) of coyotes in mixed agricultural-residential habitat while house cat (Felis catus) and squirrel (Sciurus spp. and Tamiasciurus spp.) were the two most abundant mammalian food items (13.1 and 7.8%, respectively) of coyotes in residential habitat. No single mammalian species made up >6.0% of the coyote diet in mixed forest-residential habitat. Coyotes in my western Washington study area rely on foods that result from human activity but those foods, particularly mammals, may change as land use patterns change.
Coyote (Canis latrans) populations have increased across eastern North America over the past few decades. In Illinois, red fox (Vulpes vulpes) populations have synchronously declined, suggesting that coyotes may be displacing red foxes. We examined winter (Jan-Feb) and summer (Jul-Aug) habitat use of sympatric coyotes and red foxes in east-central Illinois, including a distinct urban fox population relatively free of interactions with coyotes. We radiomarked 28 coyotes, 16 rural foxes, and 19 urban foxes and systematically collected over 10,500 locations to infer habitat use. Compositional analysis at 3 levels (home range, location, resting) corresponded to 2 spatial scales of habitat use (study area and within home-range use). We used covariate analysis of regression models to examine interspecific differences in habitat use. Using Akaike's Information Criterion (AIC), optimal models included season, sex, and species of the covariate as sources of variation. Habitat partitioning was apparent at all levels of analysis during both seasons, diverging greatly during winter. Coyotes selected and rural foxes avoided cover-rich habitats (grassland, waterways, no-till corn). Rural foxes selected human-associated habitats (active and abandoned farmsteads and rural residential areas), which coyotes generally avoided. Habitat use and home-range selection by urban foxes were more seasonally stable than by rural foxes, but urban foxes selected residential areas more during winter than during summer. Home ranges of both coyotes and rural foxes increased substantially during winter. Rural fox home ranges were nearly 4 times larger than those of urban foxes during winter. Our study demonstrates that coyotes and sympatric red foxes partition habitat seasonally in response to a highly disturbed agricultural landscape. Farmland red foxes may avoid habitats used by sympatric coyotes, relying on human-associated habitats (farmsteads and urban areas) as refugia.