Avian Scavenger Succession at Ungulate Carcasses in Washington State

by

Olivia M. Cavalluzzi

A Senior Thesis Submitted in Partial Fulfillment

of the Requirements of the Degree of

Bachelor of Science
(Wildlife Conservation)

School of Environmental and Forest Sciences

University of Washington
Box 352100
Seattle, Washington 98195-2100

2023

Approved by ______________________________________
Faculty Thesis Supervisor

Date_______________________
Abstract
Avian scavengers perform a critical ecosystem service by breaking down dead and
decomposing organic matter and recycling nutrients. However, the scavenging dynamics of birds
have received relatively little attention compared to the dynamics of scavenging mammals.
Carcasses provide an ephemeral food source for avian scavengers, which presents them with the
unique challenge of locating the carrion. This makes detection and consumption efficiency a
topic of interest as not all avian scavengers follow the same pattern of arrival at a carcass. Here,
we examine avian scavenger succession – the order in which species arrive and consume
carcasses – through monitoring ungulate carcasses in Washington State. We found that turkey
vultures outcompeted all other avian scavengers when they were present in summer months,
while black-billed magpies and common ravens outcompeted golden and bald eagles in winter
months. This confirms previous studies that have found that turkey vultures have morphological
and genetic advantages that enable them to readily find carcasses and that body size does not
determine succession in this scavenging guild. Our study highlights how social scavengers
outcompete solitary scavengers and reach carcasses sooner, regardless of the physical
characteristics of a species.

Introduction
Carcasses provide a high-quality food source for bacteria, arthropods, and a variety of
vertebrate scavengers – organisms that consume carrion – worldwide (Cortés-Avizanda et al.
2014; DeVault et al. 2003). However, the importance of carcasses to food webs and species
interactions has only recently gained attention in the scientific community (Cortés-Avizanda et
al. 2014; Barton et al. 2019). Scavengers provide a critical ecosystem service by ingesting and
recycling the remains of other animals. Without vertebrate scavengers, carcasses would persist
for longer periods of time and become a potential vector for diseases (Sekercioglu 2006;
Cunningham et al. 2018; Sonawane et al. 2021). Although many vertebrates exhibit scavenging
behavior, birds scavenge far more frequently than mammals do (Ruxton and Houston 2004,
Selva and Fortuna 2007). Mateo-Tomás et al. 2015 suggest avian species scavenge twice as
frequently as terrestrial scavengers. Vultures (Accipitridae and Cathartidae) are the only known
obligate scavengers and have distinct morphological features that make them especially well-
suited for scavenging, but many other bird species are facultative (noncompulsory) scavengers
(Sekercioglu 2006; Ogada et al. 2012).
When choosing to hunt or scavenge, animals theoretically choose the option that results
in the highest net energy reward, a concept illustrated by Optimal Foraging Theory (OFT). OFT
predicts that animals aim to get the highest energy pay-off at the lowest cost when foraging
(Brown 1988). Foraging costs include energetic costs associated with competing for or defending
a carcass and the risk of being killed by dominant predators. The value of a carcass site for
dominant predators includes both the carrion and the potential prey species scavenging on the
carrion. The Ecology of Fear, or the concept that the fear of predation has behavioral and
psychological impacts on prey species, exemplifies this concept and illustrates that magpies, a
potential prey species in this scavenging guild, may fear potential predators, such as eagles, at
carcass sites (Brown et al. 1999). OFT gives the potential for different interactions between
organisms depending on the ecosystem. Although carcasses are a largely stochastic food source
in areas with limited human influence, roads offer a consistent supply of roadkill carcasses that
avian scavengers often take advantage of, making it an ideal location to investigate an avian
scavenging guild (Fielding et al. 2021).
Despite the importance of avian scavengers, interactions between avian scavengers and
changes in carrion acquisition have not been well studied. Our study area includes one obligate
avian scavenger, turkey vulture (Cathartes aura, hereafter “vulture”), and four primary
facultative avian scavengers: common raven (Corvus corax, hereafter “raven”), black-billed
magpies (Pica hudsonia, hereafter “magpie”), golden eagle (Aquila chrysaetos), and bald eagles
(Haliaeetus leucocephalus). In this study, we ask: How does the quantity of biomass remaining
on an ungulate carcass influence carrion acquisition by and interactions between avian
scavengers? The five scavengers included in this study vary in terms of their size, morphology,
and social behavior. Golden eagles (2.4 to 6.5 kg) and bald eagles (3.0-6.3 kg) are the largest
scavengers, followed by vultures (2 kg), ravens (0.7-1.6 kg), and lastly magpies (0.1-0.2kg)
(Buehler 2022; Boarman and Heinrich 2020; Trost 2020; Katzner et al. 2020; Kirk and Mossman
2020). However, golden eagles, and occasionally bald eagles, are usually solitary scavengers
while the other three species commonly engage in social scavenging (Eden 1989, Buehler 2022,
Jackson et al. 2008, Boarman and Heinrich 2020, Trost 2020). Vultures are the only migratory
species present in our study and typically are not found during the winter months in Washington
State (Kirk and Mossman 2020).
Based on OFT, we have three hypotheses for the comparative succession of our four
target species at ungulate carcasses:
H1: Interference competition (Fig 1, A): Interference competition is a direct form of
competition where larger animals often dominate and have advantageous competitive qualities,
enabling them to restrict access to a resource, regardless of resource availability (McCormick &
Weaver 2012; Le Bourlot et al. 2014). Dominant avian scavengers (i.e. vultures, ravens, bald
eagle, golden eagle) have large body size, large search radii (able to fly great distances) and can
locate carcasses efficiently (Heinrich 1988; Wilmers et al. 2003; Katzner et al. 2020; Kirk and
Mossman 2020; Buehler 2022). Past studies have found that body size of the scavenger
determines dominance at carcass sites (Petrides 1959; Kendall 2012). Vultures are commonly the
first to carcass sights because of their olfactory senses and ability to fly low to the ground to
smell carcasses (Kirk and Mossman 2020). These large, dangerous birds may exclude more
vulnerable birds from fresh carcasses until they have been satiated. Small avian scavengers may
then arrive and forage until the carcass has been consumed. Magpies have been known to wait
until a larger predator has finished eating before consuming a carcass (Trost 2020). Interference
competition is most common when resources are abundant, concentrated, and predictable
(Maurer 1984).
H2: Exploitation Competition (Fig 1, B): Exploitation competition is an indirect form of
competition where dominant species obstruct shared, but limited, resources (Tallian et al. 2022).
Exploitation is most common when resources become concentrated, fleeting, and dispersed
throughout a landscape because it becomes advantageous to spend time foraging instead of
guarding many small and temporal food sources (Maurer 1984). Carcasses may not provide
enough carrion to counteract the energetic costs of active interference, and dominant avian
scavengers may not have the energy to defend the carcass, allowing small birds (i.e., magpies) to
forage alongside them from the beginning (Maurer 1984). Both large and small avian scavengers
would arrive at fresh carcasses and leave when they are sated as there is no reason for smaller
scavengers to fear aggression (Brown et al. 1999). Large avian scavengers would likely leave
quickly as the decreasing amounts of carrion have a lower relative energy value and would
provide suboptimal foraging returns (Le Bourlot et al. 2014). Smaller scavengers typically are at
an advantage in exploitation competition because their smaller size implies decreased food intake
and persist longer (Le Bourlot et al. 2014).
H3: Sociality (Fig 1, C): Intraspecific communication in social species (i.e. common
ravens, magpies, and turkey vultures) helps them to locate carcasses quickly while solitary
scavengers (i.e. bald and golden eagles) arrive later. Social species have been shown to dominate
carcasses (Cortés-Avizanda et al. 2014; Naves-Alegre et al. 2022). Social dominance becomes
particularly strong for unpredictable food sources where individuals can alert conspecifics to
carrion (Deygout et al. 2010).

Figure 1: Predicted outcomes for the (A) interference competition, (B) exploitation competition,
and (C) sociality hypotheses.
Methods
(a) Study area
The fieldwork for this study was carried out in inland Washington state by the
Washington Predator-Prey Project (WPPP). Our data comes from two main study areas:
Okanogan and Northwest study areas (Fig. 2). The Okanogan study area is centered around
the town of Winthrop, WA. The northeast study area is centered around the town of
Chewelah, WA. Each study area is approximately 5,000 square kilometers in size and
consists of both public and private lands. Lower elevations consist mostly of agricultural land
while montane conifer forests make up the majority of land in higher elevations. This region
of Washington is distinguished by cold, snowy winters and hot, dry summers. For the
purposes of this study, we defined winter as October through March and summer as April
through September (Kirk and Mossman 2020).

Figure 2: Northeast and Okanogan study sites in Washington State The Northeast study
area is comprised of Washington State Game management units (GMUs) Huckleberry (GMU
#121) and 49 Degrees North (GMU #117). The Okanogan study area is comprised of GMUs
Diablo (GMU #426), Pasayten (GMU #203), Chewuch (GMU #218), Gardner (GMU #231), and
Pearrygin (GMU #224).

(b) Study design and variables

WPPP technicians installed a single Reconyx (Model HyperFire 2 Pro) trail camera in
front of 50 roadkill mule deer and white-tailed deer carcasses. Field crews found recent
roadkill carcasses opportunistically and moved the carcasses to publicly owned lands less
than a kilometer from where they were originally found. Carcasses were between 3 and 226
meters from the nearest public road or highway, with a median of 29 meters. Technicians
periodically monitored sites to ensure that the camera was still monitoring focused on the
carcass. Forty-four of the carcasses were placed during summer months, and six were placed
during winter months.
The cameras were programmed to take a photograph when they sensed movement. We
tagged the species present in each photo, totaling ~37,000 images of birds, using the software
program Timelapse2. All tagging followed WPPP methodology, including golden and bald
eagles being categorized together as “eagles.” We identified the species present in each
photo, as well as the presence or absence of a carcass (Fig 3). Time and date were recorded
automatically. We bound the data from all of the cameras together into one data set where
there was a row for each species on each day of camera operation, with a column for the total
number of photos take of that species on that day and camera. We included days where zero
photos were taken so that every day of camera operation was accounted for. We only
included photos with a time interval between them of five minutes or greater so we could
model activity instead of abundance and construct an interval of foraging time. Additionally,
we added a column computing the arrival order of each species at each carcass (i.e. first
through fourth).
We are using the number of photos of each species as a proxy for carrion acquisition and
number of days since carcass deployment as a proxy for biomass remaining. The number of
photos per day will be used as a proxy for avian scavenger activity.

(c) Statistical analysis

Because the avian scavenging guild changes between seasons, we analyzed the summer
and winter seasons separately for all statistical analyses.
We modeled the daily number of counts in response to: (1) carcass age (scaled), (2)
species, and (3) season. Carcass age was the elapsed time since carcass deployment in days.
We scaled carcass age by subtracting the mean and dividing by the standard deviation to
provide numerical stability and allow our model to converge. We included a variable for
squared carcass age to best accommodate the parabolic shape of detection as carcass age
increased. We also included interaction terms between season, carcass age, carcass age
squared, and species. We included a random intercept of species nested within carcass ID to
account for nonindependence of detections on successive days at a given carcass and allow
each species its own baseline detection. Season was “summer” or “winter.”
Our main focus was to determine the composition of avian scavengers present at a
carcass as the biomass of the carcass decreased. To determine if there was a statistically
significant association between species and the order of arrival, we ran Fisher Exact Tests
(using the function fisher.test that is native to the R environment). We opted for a Fisher
Exact Test over a chi-squared test because our data violated the assumption that all cells
should have an expected count above zero.
To model how relative activity changed through time, we fit a generalized linear mixed
effect model (GLMM) to our data using a Poisson regression with a logarithm link function.
We fitted the model with Poisson and negative binomial regression and tested for over- and
under-dispersion (using function testDispersion in the package DHARMa). These tests
indicated under-dispersion, so we proceeded with the Poisson model. Because our data is
 under-dispersed, our p-values are likely conservative. We used an alpha of 0.05 to determine
the significance of all statistical tests.
We conducted all of our analyses in the R programming environment. We fitted our
GLMM using the glmer function in the package lme4.

Results
In both the summer and winter months, we found the association between species and the
order of arrival to be statistically significant. The Fisher’s Exact Test returned strong evidence
that the order of arrival was not uniform in summer (p=0.0068) or winter (p=0.00137).
Eagles were the only species that never arrived first at a carcass (Fig 3). In the summer,
vultures were the first to reach a carcass the majority of the time, followed by ravens, and to a
much lesser extent magpies. Vultures still occasionally arrived at carcasses in the winter but
never arrived first. During winter months, magpies became the first to arrive at carcasses over
half of the time, replacing vultures, who were most often first in summer months.
Each species arrived at different stages in carcass decay. In summer months, vultures
arrived in abundance before their presence tapered off (Fig 4a). Ravens and magpies typically
arrived quickly afterward and tapered off as vulture detections declined. Eagles peaked in
abundance much later in carcass decay, when other species were already decreasing in numbers.
Carcasses persisted about twice as long in the winter, and there were shifts in scavenging
succession (Fig 4b.). Vultures no longer dominated the carcass site and were few in number.
They arrived after the majority of other avian scavengers had left the carcasses. Instead, ravens
and magpies peaked in abundance early on in the carcass lifetime and gradually tapered off.
Eagles again followed behind, peaking when the magpies and ravens were decreasing in
numbers. Vultures only arrived after nearly all other avian scavengers had left.
We selected our top model because it had the lowest AICc value and was supported by
100% of model weight (table 1). The best model (table 2) included an interaction between
species, carcass age, and season, as well as an interaction between species, season, and carcass
age squared.
Our best model showed that the expected number of birds is dependent on the species and
varies across seasons. All species peak earlier in summer than winter months (Fig. 5). Vultures
and ravens are almost exclusively the predicted only avian scavengers during the summer, far in
a way outnumbering magpies and eagles, with vultures being the most dominant. All species
peak later in the winter. The predicted number of vultures and ravens decrease in winter months,
while magpies and eagles are predicted to increase. Ravens become the dominant species,
followed closely in abundance by magpies. Eagles are still relatively limited in number, but have
a more notable presence. Vultures are almost nonexistent in winter months, and peak far later
than the other species.
Figure 3: Bar plots representing the percentage of times each species reached the carcass in
different placements (first through fourth placement) during (A) summer months and (B) winter
months.
Figure 4: Plots showing the frequency for which each of the focal species arrived at different
times since technicians deployed the carcass (measured in days) during (A) summer months and
(B) winter months. The y-axis represents the number the average number of images per carcass
on a given day since carcass deployment. Scales are kept consistent across seasons but vary by
species so the relative numbers of a species between seasons is clear.
Figure 5: Predicted image count from our best mixed-effects Poisson model showing the
varying relative abundance of species across seasons.

Table 1: Model selection table for the effects of species, season, and the age of an ungulate
carcass on the number of birds expected to arrive.
Model df logLik AICc ΔAICc weight
Species*carcassAge*Season+ 25 -1869.740 3790.3 0.00 1
Season*Species*I(carcassAge^2)
Species*carcassAge*Season+ 19 -1897.899 3834.3 43.98 0
Season*I(carcassAge^2)
Species*carcassAge*Season+ 21 -1922.143 3886.9 96.57 0
Species*I(carcassAge^2)
Species*carcassAge*Season+ 18 -1973.622 3983.7 193.38 0
I(carcassAge)
Species*Season*carcassAge 17 -2277.575 4589.5 799.24 0
Species+Season*carcassAge 8 -2338.320 4692.7 902.43 0
Species*carcassAge 9 -2343.348 4704.8 914.51 0
Species*carcassAge+Season 10 -2342.976 4706.1 915.79 0
Species*Season +carcassAge 10 -2360.157 4740.5 950.15 0
 Season*carcassAge 5 -2367.541 4745.1 954.82 0
Species+carcassAge 6 -2370.774 4753.6 963.30 0
Season+carcassAge+Species 7 -2370.325 4754.7 964.42 0
carcassAge 3 -2400.268 4806.6 1016.25 0
Season+carcassAge 4 -2400.069 4808.2 1017.86 0
Species*Season 9 -2448.575 4915.3 1124.96 0
Species 5 -2458.615 4927.3 1136.97 0
Species+Season 6 -2458.531 4929.1 1138.82 0
Null 2 -2487.930 4979.9 1189.57 0
Season 3 -2487.908 4981.8 1191.53 0

Table 2: Coefficient Estimates for the most parsimonious model. Results of the final model.
Significance is based on the following codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’
Parameter Est. SE p-value
Intercept -0.299 0.512 0.560
SpeciesEagle -7.498 3.256 0.021*
SpeciesMagpie -8.911 1.455 8.97e-10***
SpeciesVulture -0.220 0.687 0.749
CarcassAge -1.996 0.298 2.19e-11***
SeasonWinter -0.2436 1.300 0.851
I(CarcassAge^2) -3.702 0.318 <2e-16***
SpeciesEagle:CarcassAge -15.750 12.832 0.220
SpeciesMagpie:CarcassAge -19.039 4.093 3.29e-06***
SpeciesVulture:CarcassAge -3.654 0.691 1.21e-07***
SpeciesEagle:SeasonWinter 5.811 3.726 0.119
SpeciesMagpie:SeasonWinter 8.700 2.251 0.000***
SpeciesVulture:SeasonWinter -7.618 2.722 0.005**
CarcassAge:SeasonWinter 1.910 0.355 7.72e-08***
SeasonWinter:I(carcassAge^2) 2.819 0.389 4.35e-13***
SpeciesEagle:I(CarcassAge^2) -18.740 12.242 0.0.126
SpeciesMagpie:I(CarcassAge^2) -15.281 3.317 4.10e-06***
SpeciesVulture: I(CarcassAge^2) -2.558 0.654 9.07e-05***
SpeciesEagle:CarcassAge: SeasonWinter 15.8448 12.836 0.217
SpeciesMagpie:CarcassAge:SeasonWinter 18.8341 4.0986 4.32e-06***
SpeciesVulture:CarcassAge:SeasonWinter 11.9722 3.700 0.001**
SpeciesEagle:SeasonWinter:I(CarcassAge^2) 17.671 12.250 0.149
SpeciesMagpie:SeasonWinter:I(CarcassAge^2) 15.042 3.328 6.19e-06***
SpeciesVulture:SeasonWinter:I(CarcassAge^2) -0.2298 2.128 0.914

Discussion
Understanding the succession of avian scavengers is fundamental to understanding the
structure and impact of a scavenging guild. Our results lend support to our sociality hypothesis,
showing how social species dominate ungulate carcass sites in an avian scavenging guild in
Washington State. Social scavengers (vultures, magpies, and ravens) were always the first to
arrive at carcass sites, with solitary scavengers (bald and golden eagles) arriving later in carcass
decomposition. Our findings suggest that communication influences the ability of a species to
find a carcass more than morphology. This is consistent with previous publications which
determined vultures, ravens, and magpies will forage socially and use intraspecific
communication to locate carrion (Eden 1989; Jackson et al. 2008; Bla ́zquez et al. 2009; Cortés-
Avizanda et al. 2014). This also aligns with previous work that found that social cues were
critical for locating unpredictable food, such as carrion (Deygout et al. 2010)
During the summer months when they are present, turkey vultures far in a way
outcompeted even the other social species for carrion acquisition. This conforms with previous
findings that vultures have morphological characteristics, physiology, and behavior that enables
them to arrive early and in large quantities at carcass sites (Cortés-Avizanda et al. 2012; Cortés-
Avizanda et al. 2014; Kane et al. 2014). In scavenging guilds in France and tropical forests,
studies have found that turkey vultures are typically the first to a carcass (Wallace & Temple
1987; Houston 1998; Potier 2019; Naves-Alegre et al. 2022).
Carcasses persisted much longer when vultures are not present in large numbers. When
vultures were largely absent, other members of the scavenging guild foraged for around twice as
many days. This is consistent with the findings of Ogada et al. 2012, who suggest that, without
vultures, carcasses will last longer, and other scavengers will stay longer. However, because
vulture absence occurs in the winter months, their absence coincides with changes in temperature
and climate, making it difficult to determine the true cause(s) of carcass persistence.
Our findings contradict previous studies which determined that size had the biggest
impact on scavenger succession (Petrides 1959; Kendall 2012). In our study, body size did not
coincide with scavenging competence, with the largest species (golden and bald eagles)
continually being the least effective members of the avian scavenging guild at locating a carcass.
Social species in our study are well suited for scavenging, regardless of their relative size
compared to other avian scavengers. Vultures have evolved particular features that prime them
for a scavenging lifestyle and easily outcompete other species. Our findings emphasize the need
for a more comprehensive study of avian scavenging guilds and the importance of avian
scavenger species to larger ecosystems.
Bird species have far-reaching impacts on ecosystems, but are typically considered
uninfluential. Vultures are the most threatened avian functional group in the world and are
declining worldwide, making it more important than ever that we access their importance
(Sekercioglu 2006; Ogada et al. 2012). Obligate and facultative scavengers have vast ecosystem
implications but are severely under-researched (Sekercioglu 2006).

Literature Cited
Barton, P.S., M.J. Evans, C.N. Foster, J.L. Pechal, J.K. Bump, M.M. Quaggiotto, and M.E.
Benbow. 2019. Towards quantifying carrion biomass in ecosystems. Trends in Ecology
and Evolution 34:950-961.
Boarman, W. I. and B. Heinrich. 2020. Common Raven (Corvus corax), version 1.0. In Birds of
the World (S. M. Billerman, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA.
https://doi-org.offcampus.lib.washington.edu/10.2173/bow.comrav.01
Brown, J. S. 1988. Patch use as an indicator of habitat preference, predation risk and
competition. Behavioral Ecology and Sociobiology 22:37–47.
Brown, J.S., Laundré, J.W. & Gurung, M. 1999. The ecology of fear: Optimal foraging, game
theory, and trophic interactions. Journal of Mammalogy 80:385-399.
Buehler, D. A. 2022. Bald Eagle (Haliaeetus leucocephalus). In Birds of the World (A. F. Poole
and F. B. Gill, Editors). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi-
org.offcampus.lib.washington.edu/10.2173/bow.baleag.01
Burnham, K.P., D.R Anderson and K.P Huyvaert. 2011. Aic model selection and multimodel
inference in behavioral ecology: Some background, observations, and comparisons.
Behavioral Ecology and Sociobiology 65:23-35.
Cortés-Avizanda, A., Jovani R., Donázar J.A., Grimm V. 2014. Bird Sky Networks: How do
avian scavengers use social information to find carrion? Journal of Ecology 95:1799–
1808.
Cunningham, C.X., C.N. Johnson, L.A. Barmuta, T. Hollings, E.J. Woehler and M.E. Jones.
2018. Top carnivore decline has cascading effects on scavengers and carrion persistence.
Proceedings of the Royal Society B: Biological Sciences 285.
DeVault T.L., Rhodes O.E., Shivik J.A. 2003. Scavenging by vertebrates: Behavioral, ecological,
and evolutionary perspectives on an important energy transfer pathway in terrestrial
ecosystems. Oikos 102:225–234.
Deygout, C., Gault, A., Duriez, O., Sarrazin, F., & Bessa-Gomes, C. 2010. Impact of food
predictability on social facilitation by foraging scavengers. Behavioral Ecology 21(6):
1131–1139
Eden S.F. 1989. The social organization of non-breeding magpies Pica Pica. Ibis 131:141–153.
Fielding, M. W., Buettel, J. C., Brook, B. W., Stojanovic, D., & Yates, L. A. 2021. Roadkill
islands: Carnivore extinction shifts seasonal use of roadside carrion by generalist avian
scavenger. The Journal of Animal Ecology 90(10): 2268–2276.
Heinrich, B. 1988. Winter foraging at carcasses by three sympatric corvids, with emphasis on
recruitment by the Raven, Corvus Corax. Behavioral Ecology and Sociobiology. 23(3):
141–156.
Houston, D. C. 1986. Scavenging efficiency of Turkey vultures in tropical forest. The Condor:
88(3):318–323.
Jackson A.L., Ruxton G.D., Houston D.C. 2008. The effect of social facilitation on foraging
success in vultures: A modelling study. Biology Letters 4:311–313.
Katzner, T. E., M. N. Kochert, K. Steenhof, C. L. McIntyre, E. H. Craig, and T. A. Miller. 2020.
Golden Eagle (Aquila chrysaetos), version 2.0. In Birds of the World (P. G. Rodewald
and B. K. Keeney, Editors). Cornell Lab of Ornithology, Ithaca, NY, USA.
Kendall, C. J. 2012. Alternative strategies in avian scavengers: How subordinate species foil the
despotic distribution. Behavioral Ecology and Sociobiology 67:383–393.
Kirk, D. A. and M. J. Mossman. 2020. Turkey Vulture (Cathartes aura), version 1.0. In Birds of
the World (A. F. Poole and F. B. Gill, Editors). Cornell Lab of Ornithology, Ithaca, NY,
USA.
Mateo-Tomás P., Olea P.P., Moleón M., Vicente J., Botella F., Selva N., Viñuela J., Sánchez-
Zapata J.A. 2015. From Regional to Global Patterns in Vertebrate Scavenger
Communities Subsidized by Big Game Hunting. Diversity and Distributions 21:913–924.
Maurer, B.A. 1984. Interference and Exploitation in Bird Communities. The Wilson Journal of
Ornithology 96:380-395.
McCormick, M. I., & Weaver, C. J. 2012. It pays to be pushy: Intracohort interference
competition between two reef fishes. PLoS ONE 7(8).
Naves-Alegre, L., Morales-Reyes, Z., Sánchez-Zapata, J. A., Sebastián-González, E., &
Ovaskainen, O. 2022. Scavenging in the realm of senses: Smell and vision drive
recruitment at carcasses in Neotropical ecosystems. Proceedings of the Royal Society B:
Biological Sciences 289(1986)
Ogada D.L, Torchin M.E., Kinnaird M.F., Ezenwa V.O. 2012. Effects of vulture declines on
facultative scavengers and potential implications for mammalian disease transmission.
Journal of Conservation Biology 26:453–460.
Petrides, G. A. 1959. Competition for food between five species of East African vultures. The
Auk 76:104–106.
Potier, S., Duriez, O., Célérier, A., Liegeois, J.-L., & Bonadonna, F. 2018. Sight or smell: Which
senses do scavenging raptors use to find food? Animal Cognition 22(1):49–59.
Ruxton G.D., Houston D.C. 2004. Obligate vertebrate scavengers must be large soaring fliers.
Journal of Theoretical Biology 228:431–436.
Sekercioglu C.H. 2006. Increasing awareness of avian ecological function. Trends in Ecology &
Evolution 21:464–471.
Selva N., Fortuna M.A.. 2007. The nested structure of a scavenger community. Proceedings of
the Royal Society B: Biological Sciences 274:1101–1108.
Sonawane, C., G. Yirga, and N. H. Carter. 2021. Public health and economic benefits of spotted
hyenas Crocuta crocuta in a peri-urban system. Journal of Applied Ecology n/a.
Tallian, A., Ordiz, A., Metz, M. C., Zimmermann, B., Wikenros, C., Smith, D. W., Stahler, D.
R., Wabakken, P., Swenson, J. E., Sand, H., & Kindberg, J. 2022. Of wolves and bears:
Seasonal drivers of interference and exploitation competition between apex predators.
Ecological Monographs: 92(2)
Trost, C. H. 2020. Black-billed Magpie (Pica hudsonia), version 1.0. In Birds of the World (S.
M. Billerman, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi-
org.offcampus.lib.washington.edu/10.2173/bow.bkbmag1.01+
Wallace, M. P., & Temple, S. A. 1987. Competitive interactions within and between species in a
guild of avian scavengers. The Auk 104(2): 290–295.
https://doi.org/10.1093/auk/104.2.290
Wilmers, C. C., R. L. Crabtree, D. W. Smith, K. M. Murphy and W. M. Getz. 2003. Trophic
facilitation by introduced top predators: Grey Wolf subsidies to scavengers in
Yellowstone National park. Journal of Animal Ecology 72:909–916.
Acknowledgments
We respectfully acknowledge that this study took place on the stolen lands of the
Kalispel, Spokane, and Colville Confederated Tribes. We conducted our research through
University of Washington, which is located on the stolen ancestral lands of the Coast Salish,
Duwamish, and Suquamish peoples, who have lived in the Salish Sea basin and throughout the
San Juan Islands and the North Cascades watershed since time immemorial.
Special thanks to Rebecca Windell, Carmen Vanbianchi, Anna Machowicz, and all of the
volunteer technicians for their contributions to the data.
Scholarship support for O. Cavalluzzi was received from the Mary Gates Endowment.