Vertebrate Scavenger Species Composition on Nantucket
Island, Massachusetts and Implications for Nicrophorus
americanus Olivier (American Burying Beetle)
Reintroduction
Andrew Mckenna-Foster, Danielle O’Dell, Elisabeth Sorrows,Jacqueline Laurenzano, and Lou Perrotti
Northeastern Naturalist, Volume 26, Issue 1 (2019): 52–62
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A. Mckenna-Foster, D. O’Dell, E. Sorrows, J. Laurenzano, and L. Perrotti
22001199 NORTHEASTERN NATURALIST V2o6l.( 12)6:,5 N2–o6. 21
Vertebrate Scavenger Species Composition on Nantucket
Island, Massachusetts and Implications for Nicrophorus
americanus Olivier (American Burying Beetle)
Reintroduction
Andrew Mckenna-Foster1,*, Danielle O’Dell2, Elisabeth Sorrows1,
Jacqueline Laurenzano1, and Lou Perrotti3
Abstract - We surveyed scavenger species on Nantucket Island, MA, using motion-activated
wildlife cameras baited with carcasses of either a Mus musculus domesticus (House Mouse)
or Colinus virginianus (Bobwhite Quail) in the summer of 2016. We aimed to identify vertebrate
scavenger species, measure how long carcasses remained available, and examine
differences in carrion consumption between grassland and shrubland vegetation types. This
information is important for management decisions concerning a reintroduced population
of a federally endangered obligate scavenger, Nicrophorus americanus (American Burying
Beetle), on Nantucket Island. Carcasses were removed or consumed within 2.7 ± 1.8 d, and
vertebrates removed the carcass in 74% of trials. Carcasses persisted for similar lengths of
time between the 2 vegetation types, but avian scavengers were significantly more common
on Bobwhite Quail carcasses in grassland, and mammalian scavengers were significantly
more common on House Mouse carcasses in shrubland habitats. Avian species appear to be
significant competitors for carrion appropriate for American Burying Beetle reproduction.
Introduction
Nantucket Island, MA, is the location of 1 of only 2 successful reintroductions
of the federally endangered beetle, Nicrophorus americanus (Olivier) (American
Burying Beetle, hereafter ABB) (Mckenna-Foster et al. 2016). The ABB is North
America’s largest carrion beetle (family Silphidae) with individuals up to 3.5 cm
in body length. As a scavenger species, the ABB requires for reproduction a vertebrate
carcass of 80–200 g, and the species is a direct competitor with vertebrate
and invertebrate scavengers (Kozol et al. 1988, Raithel 2000). A beetle must find a
carcass before it has been degraded or removed by other scavengers, attract a mate,
and then bury the carcass in the soil. In one of the few examples of bi-parental
care among non-social terrestrial insects, the beetle pair then rears up to 30 larvae
on the carcass (Raithel 2000). The US Fish and Wildlife Service reintroduced
captive-reared ABBs to Nantucket Island each summer between 1994 and 2006,
and assisted the growing population in the following years through a regime of
trapping in situ individuals, pairing them, and, in a process called provisioning,
burying these pairs with vertebrate carcasses near the trap sites (Mckenna-Foster
1Maria Mitchell Association, 4 Vestal Street, Nantucket, MA 02554. 2Nantucket Conservation
Foundation, PO Box 13, Nantucket, MA 02554. 3Roger Williams Park Zoo, 1000 Elmwood
Avenue, Providence, RI 02907. *Corresponding author - andrew.mckennafoster@gmail.com.
Manuscript Editor: Christopher M. Heckscher
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et al. 2016). This project is considered to have been successful because the trapping
and provisioning methods appeared to increase the ABB population each year
(Mckenna-Foster et al. 2016). However, carrion provisioning was reduced by 75%
in 2011 and the population began a steady decline through 2016, which raised
the question of whether ABBs are able to compete with Nantucket scavengers for
naturally available carrion (Mckenna-Foster et al. 2016). The ultimate goal is to
establish a self-sustaining ABB population on Nantucket; thus, it is critical that
managers understand the scavenger community on this isolated island and its impacts
on carrion availability.
Scavengers and the vertebrate carrion they consume significantly affect nutrient
cycling (Wilson and Wolkovich 2011), community structure (e.g., Vucetich et
al. 2004), and local biodiversity (Barton et al. 2013). Scavenging systems remain
understudied, especially in the case of small vertebrate carrion (Procyon lotor
(L.) [Raccoon]-sized or smaller), due to the difficulty of censusing natural carrion
resources, complex interactions between facultative and obligate scavengers, and
variability in scavenger composition across vegetation communities and seasons
(Beasley et al. 2016, DeVault et al. 2003, Turner et al. 2017). In the last 2 centuries,
loss of habitat, reductions of apex predators, increases in mesopredator populations,
species extinction, and species invasion have changed the composition of vertebrate
scavenger species and carrion resources in North America (Beasley et al. 2016,
Prugh et al. 2009). Habitat fragmentation has helped 60 percent of North American
mesopredator species expand their ranges (Prugh et al. 2009), and these species
decrease local populations of small prey (Crooks and Soulé 1999) and monopolize
carrion resources (DeVault et al. 2011). These changes are assumed to be partly
responsible for a 90% reduction in the range and population of the ABB starting
in the early 20th century (Sikes and Raithel 2002). Today, the ABB faces intense
competition from vertebrate scavengers. In North America, vertebrate scavenging
efficiency (percent of available carcasses utilized by vertebrate scavengers) averages
75% and can be up to 90% (DeVault et al. 2003). In agricultural landscapes,
small carrion is usually consumed by vertebrates within 3 d (DeVault et al. 2004),
and much of the scavenging is dominated by mesopredator species like Raccoons
and Didelphis virginiana (Kerr) (Virginia Opossum) (DeVault et al. 2011). Persistence
time may be longer in natural landscapes. Turner et al. (2017) recorded Rattus
spp. (rat) carcasses persisting an average of 3.3 d and DeVault et al. (2017) recorded
Colinus spp. (quail)-sized bird carcasses persisting for 7–9 d. Finally, Conley et
al. (2015) found some evidence that mesopredators are more dominant scavengers
in forested sites and at night, while avian species are more common in grasslands
and during the day. Those authors suggested that there is both temporal and habitat
partitioning between the 2 groups.
Nantucket was chosen as a reintroduction site for the ABB as part of a federal
recovery plan partly because the island lacks common North American mammalian
mesopredators, including Raccoons, Virginia opossum, Vulpes vulpes (L.) (Red
Fox), Urocyon cinereorgenteus (Schreber) (Gray Fox), and Mephitis mephitis
(Schreber) (Striped Skunk), all of which are assumed competitors of the ABB on the
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2019 Vol. 26, No. 1
mainland (DeVault et al. 2011, Sikes and Raithel 2002). It is believed that Native
American and early European inhabitants of the island extirpated these mesopredators
(Zube and Carlozzi 1967), though since that time, both Felis silvestris catus L.
(Domestic Cat) and Rattus norvegicus (Berkenhout) (Norway Rat) have established
populations on the island. As a reproductively obligate scavenger species, the ABB
competes for carrion resources with invertebrate, mammalian, and avian scavengers,
and it is difficult to predict what conditions help the ABB succeed.
A growing body of literature on scavenging ecology from natural and agricultural
settings suggest that the fate of a carcass and the scavenger species utilizing
that carcass are influenced by a combination of biotic and abiotic factors. Higher
temperatures inhibit vertebrate scavenger species and promote invertebrate scavenging
(Beasley et al. 2016, DeVault et al. 2011), and micro-site variation in
temperature and humidity may significantly affect the rate of carcass decomposition
(e.g., Parmenter and MacMahon 2009). The type of carcass influences how
long it persists and which scavengers will consume it; predator carcasses are consumed
at a lower rate than other vertebrates (DeVault et al. 2017), and scavengers
may avoid conspecific carcasses (Olson et al. 2016). Carcass size affects both
scavenger community composition and carcass longevity. Smaller carcasses attract
smaller mesopredators and persist for a shorter amount of time (Turner et al. 2017).
Both Turner et al. (2017) and Abernethy et al. (2016) provide evidence that habitat
type interacts with carcass type and scavenger species to influence carcass detection
and longevity. Turner et al. (2017) found that scavengers detected Sus sp. (pig)
carcasses significantly faster than smaller carcasses in a clear-cut habitat compared
to forested habitat. However, smaller carcasses (e.g., rat and rabbit) were detected
at the same rates across habitats.
Combining what is known about scavenging ecology with scavenger data from
Nantucket could inform new management strategies to establish a self-sustaining
ABB population on the island. Understanding the fate of a carcass on the island is
a first step in this process. Managers need site-specific information on scavenger
composition and scavenger efficiency from vegetation types utilized by the ABB.
The goals of this study were to (1) understand which scavengers were competing
with ABB in 2 common vegetation types, and (2) determine carcass persistence
and vertebrate scavenging efficiency for a carcass suitable for use by the ABB in
2 common vegetation types. Nantucket lacks most mesopredators; thus, we hypothesized
that avian species would dominate the scavenger community. Based
on other studies, we predicted Corvus brachyrhynchos (American Crow), Cathartes
aura (Turkey Vulture), and Cyanocitta cristata (L.) (Blue Jay) would be
common visitors to carrion (DeVault et al. 2004, 2017; Olson et al. 2016). We also
hypothesized that carrion in shrubby vegetation would be more difficult to detect
and access and therefore persist longer than in grassland. There are no North
American studies of systems that contain common avian scavengers but not the
common mesopredators, so it was difficult to predict how a lack of mesopredators
would affect carcass persistence or scavenging efficiency. Olson et al. (2016)
saw avian species feeding on 60% of medium-sized carcasses (rabbit, Virginia
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Opossum, or Raccoon), while DeVault et al. (2011) recorded avians scavenging
only 8% of mouse or rat carcasses. Avians are regularly observed visiting carrion,
and Nantucket has populations of many of the common avian scavengers; thus, we
predicted carcass persistence and scavenging efficiency similar to mainland observations
for small carrion.
Field-site Description
Nantucket Island lies 50 km south of Cape Cod, MA, and contains ~130 km2 of
sandplain grassland, coastal heathland, barrier beach, and mixed forest (Fig. 1). We
selected camera sites within large tracts of conservation land that protect diverse
sandplain grassland and coastal shrubland vegetation communities. Quercus ilicifolia
Wangenheim (Scrub Oak) and Pinus rigida Miller (Pitch Pine) grow in small
patches among a mosaic of low grasses and shrubs. The ABB range encompasses
the eastern half of the island, where these 2 vegetation communities are common
(Mckenna-Foster et al. 2016) (Fig. 1).
Figure 1. The distribution of camera sites on Nantucket Island, MA. We deployed 2 cameras
at each sample site to identify scavenger species. Triangles represent sample sites and are
not to scale. The dark boundary outlines the assumed range of Nicrophorus americanus
(Olivier) (American Burying Beetle).
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Methods
Between 22 June and 3 August 2016, we deployed 18 Browning BTC-5 wildlife
cameras (Browning Trail Cameras, Birmingham, AL) in pairs at 9 sites across the island,
and baited the camera stations weekly with either a deceased, dark-haired Mus
musculus domesticus L. (House Mouse [hereafter Mouse], average mass = 32.4 g)
or a Colinus virginianus (L.) (Bobwhite Quail [hereafter, Quail]; average mass =
132.6 g) (Fig. 1). We used Quail during the first 3 weeks to coincide with peak ABB
reproductive activity and Mice the last 3 weeks to extend the study duration. While
this change in carrion type and size likely affected which scavengers visited our
sites, the focus of our study was a general species census for small-carrion scavengers
rather than a determination of scavenger preferences for different carrion types.
ABBs will feed on any available carrion, and Mouse-sized carcasses are a potential
food resource (Raithel 2000). We placed all 6 of the eastern camera-trap sites near
well-established ABB trap lines. We also placed camera-trap sites in potential ABB
habitat on the west side of the island. At each site, we placed 1 camera in a grassland
and 1 camera in closed-canopy shrubland, ~100 m apart. The cameras were generally
within 10 m of a dirt road. We did not record temperature at each site, but as a
proxy, report average temperatures recorded at the Nantucket Memorial Airport and
retrieved from the National Oceanic and Atmospheric Association (NOAA 2018).
Data from temperature logging iButtons (Maxim Integrated, San Jose, CA), placed
at 5 of our camera-trap sites in June 2015, showed that the difference in ground temperature
between the airport and our sample sites was less than 2 °C. The ground temperature
at these sites was 5.1 ± 2.9 °C hotter in the daytime and 1.7 ± 1.4 °C cooler in the
nighttime than the air temperature at the Nantucket airport.
We constructed wooden stakes to hold the camera over the carrion in the grassland
sites. In the shrubland sites, we attached cameras to sturdy branches or used a
wood stake when necessary. All cameras faced the ground from a height of 1–2 m.
We secured the bait to a turf staple directly below the camera using dental floss
tied to a rear leg to prevent the carcass from being carried out of the view of the
camera. We replaced the bait and swapped out SD cards and camera batteries every
7 d. We also moved the cameras to a new location at least 100 m away from the
previous week’s location to reduce habituation to a static bait-station and more accurately
mimic natural carrion availability. The cameras took 2 pictures per trigger
event and waited 10 secs before resetting. Before placing the carcass underneath
the camera, we trimmed the grass and small shrubs to reduce the chance of windblown
vegetation triggering the camera.
We identified any animal seen in each picture frame, noted general behavior
(moving through picture frame, standing, investigating carcass, actively consuming
carcass), location of the animal in the picture frame in relation to the carcass,
and the condition of the carcass. Small rodents visiting carcasses could be difficult
to identify, especially in nighttime photographs, and when a positive identification
could not be made, we recorded them under the general classification of “rodent”.
If the same species visited a carcass within 5 min, we assumed it was the same
individual and recorded it as a single scavenging event. We considered a carcass
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to be consumed if it was either removed or if only bones, fur, or feathers remained.
This decomposition stage was relatively easy to identify, and a carcass that reached
this stage was generally left undisturbed by scavengers.
Results
The cameras took 69,835 photographs of which 2% contained a vertebrate or
invertebrate species. Most photographs were triggered by moving vegetation or
an unidentifiable trigger. Average daily temperature on the island each week in
the study period was 20 ± 3 °C and varied from 18 to 23 °C (NOAA 2018). Due
to equipment failures, we collected data from 87 of the 108 possible trials. A total
of 43 trials used Quail carrion and 44 trials used Mice. We were able to identify
which scavenger consumed the carcass in 65 trials (Table 1). In 19 trials, the camera
did not photograph the scavenger that removed the carcass; in 2 trials, the carrion
disintegrated as a result of general scavenging by vertebrate and invertebrate species;
and in 1 trial the scavengers did not visit the Mouse carcass and it remained
relatively intact the entire week. Vertebrates consumed or removed carrion in 74%
Table 1. Species that removed Mouse or Quail carcasses on Nantucket Island, MA. We classified carcasses
that slowly degraded over time due to multiple scavengers and cases where the camera failed
to capture the scavenger as “Unknown”. The “Carrion beetles” classification represents cases where a
variety of species in the families Scarabaeidae and Silphidae consumed the carcass. We used the “Unidentified
insects” classification when a combination of flies, an ts, and beetles consumed the carrion.
Scavenger Quail Mouse Total
Mammals (35%)
Unidentified rodents 4 6 10 (15%)
Peromyscus leucopus (Rafinesque) (White-footed Mouse) 3 5 8 (12%)
Rattus sp. (rat) 0 2 2 (3%)
Canis lupus familiaris L. (Dog) 1 0 1 (2%)
Blarina brevicauda (Say) Short-tailed Shrew 0 1 1 (2%)
Felis silvestris catus (L.) (Domestic Cat) 0 1 1 (2%)
Avians (39%)
Corvus brachyrhynchos Brehm (American Crow) 7 4 11 (17%)
Buteo jamaicensis (Gmelin) (Red-tailed Hawk) 5 2 7 (11%)
Cathartes aura (L.) (Turkey Vulture) 6 0 6 (9%)
Circus hudsonius (L.) (Northern Harrier) 0 1 1 (2%)
Invertebrates (26%)
Diptera spp. (flies) 3 2 5 (8%)
Unidentified insects 1 2 3 (5%)
Formicidae spp. (ants) 0 3 3 (5%)
Nicrophorus spp. (burying beetles) 1 2 3 (5%)
Silphidae spp. (carrion beetles) 2 0 2 (5%)
Nicrophorus marginatus Fabricius (Margined Burying Beetle) 0 1 1 (2%)
Subtotal 33 32 65
Unknown 10 9 19
Not scavenged 0 1 1
Multiple species 0 2 2
Total 43 44 87
Camera failed 12 9 21
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of trials where we could identify the scavenger (48/65). It took an average of 2.7
± 1.8 d for the carrion to be consumed in place or removed. There was no significant
difference in the number of days it took scavengers to consume a carcass in
shrubland or grassland sites (unpaired t-test: df = 78, P = 0.44). In 53% of trials,
scavengers first made contact with a carcass within 12 h, and 92% of carcasses had
been located within 36 h. The surrounding vegetation did not significantly influence
the time to discovery (unpaired t-test: df = 86, P = 0.10).
We identified 9 vertebrate species removing carrion (Table 1). Five species or
species groups accounted for 65% of scavenging events and all are species seen in
mainland scavenger studies: American Crows, unidentified rodents, Peromyscus
leucopus (White-footed Mouse), Buteo jamaicensis (Red-tailed Hawk), and Turkey
Vultures. Many of the unidentified rodents were most likely White-footed Mice. We
also observed species not often reported in other studies including Circus hudsonius
(Northern Harrier) and Norway Rats. Domestic Cats visited 3 Quail carcasses,
2 in grassland and 1 in shrubland habitat, and removed 1 Mouse carcass from a
grassland site.
We compared the number of carcasses removed by avians, mammals, or invertebrates
overall and within each combination of carcass type and vegetation type
(Fig. 2). Due to small sample sizes, we used the exact multinomial goodness-offit
test in all comparisons (XNomial package; R Core Team 2013). There was no
significant difference in the overall number of carcasses removed by avians (39%),
mammals (35%), or invertebrates (26%) (P = 0.45). The proportion of carcasses
removed by these 3 scavenger groups differed significantly for Quail carcasses in
grassland and Mouse carcasses in shrubland (P = 0.01 and P = 0.01, respectively).
Within these 2 categories, we performed a post hoc analysis using the exact binomial
goodness-of-fit test with an adjusted alpha level of 0.02 (0.05/3). Avians scavenged
significantly more Quail carcasses in grassland than expected (P = 0.002), while
mammals scavenged significantly more Mouse carcasses in shrubland (P = 0.002).
Overall, avian species removed a majority of Quail carcasses (55%) and mammals
removed the largest percentage of Mouse carcasses (44%) though neither was significant
(Quail: P = 0.05, mouse: P = 0.25).
No ABBs visited carcasses, although related species of Nicrophorus carrion
beetles buried 3 Mouse carcasses and 1 Quail carcass (4.5% total). Nicrophorus
marginatus (Margined Burying Beetle) buried 1 Mouse but we could only narrow
the other burying events to either N. marginatus or N. orbicollis Say (Roundneck
Sexton Beetle). Invertebrates consumed another 13 carcasses. In 10 of these cases,
we could clearly identify flies, ants, or carrion beetles (representing a variety of
species in the families Scarabaeidae and Silphidae) as the primary scavenger and
ultimate consumer of a carcass.
Discussion
The scavenging community on Nantucket is comprised of avians, rodents,
and invertebrates. In the absence of mammalian mesopredators, rodents and
invertebrates consume a larger percentage of carcasses than reported elsewhere.
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DeVault et al. (2017) recorded avians and mesopredators scavenging 28% and
67% of carcasses respectively, while Olson et al. (2016) recorded 41% and 57%
respectively. Neither study observed rodents or invertebrates as significant scavengers.
In our study, avians, rodents, and invertebrates each consumed about one
third of the carcasses.
Our findings support the hypotheses that small carrion on Nantucket would both
be scavenged primarily by vertebrates and at a similar rate as mainland carrion. Vertebrates
scavenged 74% of carrion and carrion persisted for about 3 d, well within
the ranges of values reported elsewhere (DeVault et al. 2003, 2004; Turner et al.
2017). Contrary to our expectations, carcasses in shrubland did not persist longer
than carcasses in grassland. This result was due to the presence of an efficient
Figure 2. Proportion of carcasses removed by avians, invertebrates, or mammals and categorized
by vegetation and carcass type. Letters identify significant differences. Avian
scavengers removed significantly more Quail carcasses in grasslands, and mammals removed
significantly more Mouse carcasses in shrublands.
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community of rodent scavengers that consumed a significant number of carcasses
in shrubland. Conversely, while we expected avian scavengers to dominate across
vegetation types, they were only dominant on Quail carcasses in grassland, perhaps
because these carcasses were easier to detect and access. Conley et al. (2015) found
similar habitat partitioning among mammalian and avian species, though the mammalian
species were all mesopredators.
Nantucket scavengers may be a more significant factor in establishing a
self-sustaining ABB population than originally thought. During the peak ABB
reproductive period in late June, avian species are the main competitor for carrion
that is suitable for ABB reproduction. The high number of Quail carcasses
scavenged by avians was largely due to Turkey Vultures, which we observed only
feeding on Quail carcasses and only during the first 2 weeks of our sampling period.
Red-tailed Hawk and American Crows were also common scavengers during
that time, removing 4 Quail carcasses each. American Crows continued to regularly
scavenge carcasses throughout the rest of the study. The ABB is reproductively
active into mid-July on Nantucket and we assume American Crows and Red-tailed
Hawks remain important consumers of Quail-sized carrion throughout this time.
However, time and carcass type are confounding variables because we did not
deploy Quail and Mice simultaneously. We cannot know from our data whether
avians scavenged more Quail carcasses because of the time of year or because
they prefer larger or non-mammal carrion.
In addition to vertebrate species, we found evidence that the ABB faces competition
from other Nicrophorus beetles. Nicrophorus species compete both in the race
to locate a carcass and in direct confrontation for a carcass. The ABB can compete
well in direct confrontation (Kozol et al. 1988) but may be outcompeted by congener
species in the search for carcasses (Matthews 1995). We attribute the lack of
ABB sightings in our study to a small population size and the recruitment of beetles
to monitoring pitfall traps set within the same study area. In June 2016, the monitoring
traps captured 24 ABBs, the lowest number in 12 y of trapping (Mckenna-Foster
et al. 2016). These traps were baited with odoriferous rotted chicken and were in
some cases within 50 m of the camera sites, potentially attracting away the limited
number of ABBs. Margined Burying Beetle and Roundneck Sexton Beetle are also
attracted to the traps, but their populations are much larger than the ABB, increasing
their chances for detection at camera sites (A. Mckenna-Foster, unpubl. data).
This survey provides important information to inform ABB management decisions.
Despite a lack of mammalian mesopredators, the ABB on Nantucket faces
similar levels of competition for its reproductive resource compared to what it
might face on the mainland. Efforts to establish a self-sustaining ABB population
will have to account for this effective scavenging community. Further work should
investigate carcass-type preference for Nantucket scavengers and ABB success at
locating carcasses in grassland and shrubland. ABBs can use a wide variety of carcass
types (mammal or avian) for reproduction if the carcass is within the correct
size range (Amaral et al. 1997, Kozol et al. 1988). It could be useful for managers
to know if some types are scavenged at a lower rate compared to others. Avians
and rodents appear to be more effective scavengers in grassland and shrubland,
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respectively; thus, understanding how the ABB uses those vegetation types on Nantucket
could guide future recovery efforts.
Acknowledgments
This project was partly funded by grant #15-233 from the Norcross Wildlife Foundation.
The Roger Williams Park Zoo, Providence, RI, generously provided Quail carcasses and
field equipment. Alex Etkind, Skyler Kardell, Corrina Marshall, and David Ryan assisted
with field work and photo processing. We thank Jennifer Karberg and 2 anonymous reviewers
for providing valuable comments.
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