Artificial Nests Identify Possible Nest Predators of Eastern
Wild Turkeys
Haemish I.A.S. Melville, Warren C. Conway, Michael L. Morrison, Christopher E. Comer, and Jason B. Hardin
Southeastern Naturalist, Volume 13, Issue 1 (2014): 80–91
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Southeastern Naturalist
H.I.A.S. Melville, W.C. Conway, M.L. Morrison, C.E. Comer, and J.B. Hardin
2014 Vol. 13, No. 1
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2014 SOUTHEASTERN NATURALIST 13(1):80–91
Artificial Nests Identify Possible Nest Predators of Eastern
Wild Turkeys
Haemish I.A.S. Melville1,2,*, Warren C. Conway3, Michael L. Morrison1,
Christopher E. Comer3, and Jason B. Hardin4
Abstract - Poor nest survival is a critical limiting factor in the recruitment of wild birds.
Nest predation is often cited as one of the main causes of nest failure, especially for
ground-nesting species. We monitored artificial Meleagris gallopavo silvestris (Eastern
Wild Turkey) nests, using time-lapse and motion-sensitive trail cameras to determine which
predators were likely to be responsible for preying on Wild Turkey nests in the Pineywoods
of East Texas. Sixty-one percent of all artificial nests were preyed upon. Corvus brachyrhynchos
(American Crow) preyed on 48% of artificial nests and Procyon lotor (Raccoon)
preyed on 35%. There was a seasonal increase in the number of artificial nests preyed upon
from spring to summer. Mammalian mesopredators, Raccoon and Didelphis virginiana
(Opossum), were primarily responsible for this increase, suggesting an increase in search effort
by mesopredators that coincided with increased dietary diversity in a period of reduced
prey resources. Predators other than mesopredators—American Crows, Picoides sp. (woodpeckers),
Dasypus novemcinctus (Nine-banded Armadillo), and snakes—were responsible
for 53% of all predation on the artificial Wild Turkey nests, with American Crows being
the most important of these. After nest deployment, American Crows located and preyed
on artificial nests more quickly than other nest predators. We suggest video monitoring of
natural Eastern Wild Turkey nests to confirm the identity of nest predators.
Introduction
The degree to which predators affect their prey resources is central to the study
of ecology (Begon et al. 2006). It is of overriding importance when considering
the control of abundant prey species, the conservation of endangered prey species
(Macdonald et al. 1999), and prey species of ecological, economic, or recreational
importance. The effects of predation can make the difference between persistence
and local extinction of rare prey species (Caughley and Sinclair 1994, Macdonald
et al. 1999). Predation may interact with other causes of mortality, and the additive
effect may cause the collapse of prey populations (Macdonald et al. 1999).
Nesting by Meleagris gallopavo silvestris Vieillot (Eastern Wild Turkey) is
confined to the spring months (late March to late June; Healy 1992, Isabelle 2010).
Poor nest survival is one of the primary limitations to the successful recruitment of
bird species (Dreibelbis et al. 2008). Predation is often cited as the main cause of
nest fatality in avian species (Martin 1993a; Mezquida 2001, 2003; Ricklefs 1969;
1Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station,
TX 77843. 2Current address - Number 8 25th Street, Zakher, Al Ain, United Arab Emirates.
3Arthur Temple College of Forestry and Agriculture, Stephen F. Austin State University,
Nacogdoches, TX 75962. Texas Parks and Wildlife Department, Oakwood, TX 75855.
*Corresponding author - haemish.melville@gmail.com.
Manuscript Editor: Roger D. Applegate
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Rotenberry 1989), and ground-nesting birds are particularly vulnerable to mammalian
and avian predation (Fletcher et al. 2010, Marcstrom et al. 1988, Newton
1993). Meleagris gallopavo L. (Wild Turkey) is a ground-nesting species of which
hens, nests, and young poults are especially vulnerable to predation (Glidden 1975,
Miller and Leopold 1992, Speake 1980). Wild Turkeys also have a long incubation
period, approximately 26 days (Healy 1992, Williams et al. 1971), which extends
its period of vulnerability. Eastern Wild Turkeys are not known to display any
defensive behaviors against mammalian predators; rather, they have evolved adaptations
including morphological and behavioral crypsis, large size, and exceptional
eyesight to counter predator pressure (Leopold and Chamberlain 2002).
In the US, Wild Turkeys were on the brink of extinction through habitat loss
and hunting (Lopez et al. 2000, Kennamer et al. 1992). Attempts to restore the
Wild Turkey have generally been successful nationwide (Kennamer et al. 1992),
but this has not been the case in the Pineywoods of East Texas, despite the translocation
of over 7000 Eastern Wild Turkeys into the state since 1979 (Boyd and
Oglesby 1975, Isabelle 2010, Seidel 2010). Despite reintroduced turkeys achieving
nest-success rates comparable to those in areas where Wild Turkeys have been
successfully reintroduced (Isabelle 2010, Vangilder 1992), no self-sustaining
population has been established.
Predation rates differ for artificial and natural nests. There is no conclusive
evidence that artificial nests and natural nests show a consistent trend or relationship
to one another in terms of predation rate (Mezquida 2003). Although
artificial nests are useful to identify potential nest predators (Wilson et al. 1998),
they can attract predators that would not usually locate or depredate live nests
(Dreibelblis et al. 2011, Willebrand and Marcstrom 1988). Differential egg characteristics
have been demonstrated to affect predation rates (Lindell 2000, Major
and Kendal 1996). Nest characteristics may vary between real and artificial nests;
for example, artificial nests are often located in habitats that birds would not select
for nesting. Generally the predation rate on artificial nests has been found to
be greater than on real nests (MacIvor et al. 1990, Major and Kendal 1996, Roper
1992, Salonen and Penttinen 1988).
Where the study species is rare, it is often not possible to find enough real nests
to assess factors that influence nest predation (Reitsma et al. 1990). In addition,
when the influence of monitoring live nests might compromise the success of nests,
it is not ethical to monitor real nests. Consequently, we concluded that, despite their
limitations, the use of artificial nests was the best means to investigate the variables
that might influence predation on Eastern Wild Turkey nests in the Pineywoods.
With the above provisos in mind, we developed an experiment to determine which
predators would prey on artificial Wild Turkey nests, whether nest sites selected
by Wild Turkeys were less susceptible to predation than sites in random locations,
whether there was differential seasonal predation on artificial nests, and whether the
presence of a proxy for the Wild Turkey hen at the nest would influence predation
on the artificial nest.
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Study Area
We conducted this study from January 2009 to September 2011 at two sites in
the heart of the Pineywoods ecoregion in Nacogdoches and Angelina counties of
East Texas: study site 1 (UTM coordinates: 337710N, 3486410W; 1360 ha) and
study site 2 (372302N, 3478315W; 5000 ha). We selected these properties because
they represent typical Wild Turkey habitat in East Texas, they both support stable
populations of Eastern Wild Turkeys, and several translocations have been attempted
in the vicinity of these properties (Isabelle 2010).
The Pineywoods occur in the western Gulf coastal region and extend through
east Texas, northwestern Louisiana, and southwestern Arkansas. Little of the
original Pinus palustris Mill. (Longleaf Pine) and Pinus echinata Mill. (Shortleaf
Pine) forests remain. They have been largely replaced by even-aged Pinus taeda L.
(Loblolly Pine) plantations. Much of the natural vegetation of the Pineywoods has
been altered due to the planting of pine and the exclusion of fire (Omernik et al.
2008). In addition, 63% of the region’s land is in private ownership, with much of
this in relatively small parcels of 0.4–3.6 ha. One consequence of small parcel size
is forest fragmentation (US Forest Service 2002), which has substantially altered
the habitat historically used by Wild Turkeys.
The mean annual rainfall in the Pineywoods is 1192 mm, with a monthly mean
that varies between a low of 55 mm in July and a maximum of 116.4 mm in May.
The mean annual minimum temperature is 12.8 °C, and the mean annual maximum
temperature 25.5 °C. The mean maximum temperature in the summer is 35 °C
(Sivanpillai 2005). During our study, the mean annual temperature was 19.4 °C, the
minimum temperature recorded was -5.3 °C, and the maximum temperature was 38
°C (NOAA 2012). The mean annual rainfall during our study was 1015 mm, with
the greatest rainfall occurring in 2009 (1243 mm) and the lowest in 2011 (832 mm)
(NOAA 2012).
Methods
We implemented a manipulative experiment using artificial Wild Turkey nests
and motion-sensing, time-lapse photography. We developed an experimental
protocol to minimize most problems associated with artificial nest studies. This
approach entailed leaving minimal human scent (by wearing latex gloves while
handling any equipment and eggs, and by ensuring that field workers wore rubber
boots and latex gloves while setting up artificial nests and positioning cameras),
not revisiting the artificial nest sites during the experiment, using a proxy for the
presence of a Wild Turkey female, adding real Wild Turkey feathers to the artificial
nest as scent cues, and using nest-sites that were known to have been used
by Wild Turkeys in the past (Isabelle 2010). Because the Gallus domesticus (L.)
(Domestic Chicken) eggs we used are smaller than Wild Turkey eggs, they did not
limit predators’ ability to consume the eggs. We avoided deploying the artificial
nests in a systematic pattern; rather, within each study site, we located the nests in
locations used previously by Wild Turkeys (see Isabelle 2010).
We conducted this experiment seasonally from spring 2009 to fall 2011. Spring
and summer coincided with the nesting season for Eastern Wild Turkeys in the
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Pineywoods (Isabelle 2010). This seasonal approach allowed us to assess whether
mesopredators searched for artificial Wild Turkey nests seasonally (e.g., greater effort
during spring when Wild Turkey nests are naturally available), or whether they
encountered the artificial Wild Turkey nests randomly as a result of their movements
in their home-ranges. We defined the seasons as: spring (21 March–20 June),
summer (21 June–20 September), fall (21 September–20 December), and winter
(21 December–20 March).
Each year, we placed artificial nests in 5 randomly selected locations at each study
site known to have been used by a Wild Turkey the year before (Isabelle 2010). Wild
Turkey nests consist of shallow depressions formed by the hen and lined with leaves
and feathers (Healy 1992). To create the artificial nests, we formed ≈20-cm-diameter
depressions in the ground with our hands and positioned leaves and Wild Turkey
feathers in these depressions to simulate the appearance of natural nests. Using GIS,
we randomly selected another 5 locations at each study site for the placement of
artificial nests. We navigated to each randomly selected location, and set-up the artificial
nest in the nearest position that approximated a typical Wild Turkey nest setting
(Isabelle 2010). We changed locations used for artificial nests each year to facilitate
comparisons between predators that preyed on artificial Wild Turkey nests located in
known nesting areas with artificial Wild Turkey nests positioned outside documented
nesting areas. After one of our cameras failed in spring 2010, we monitored 19 artificial
nests for the remainder of the study.
We deployed the artificial nests and then left them unchecked for 14 days (half
the Wild Turkey incubation period; Campo et al. 1989, Isabelle 2010, Ransom et al.
1987). We did not leave the eggs in place for >14 days because if the eggs rotted,
their odor would change the olfactory signature. We did not replace eggs during
the 14-d period because our visit might have compromised the nests by leaving additional
human olfactory and visual cues for predators.
We positioned decoys (Feather Flex® 3-position Hen Turkey Decoys) representing
sitting Wild Turkey females on 5 of the artificial nests located at the documented
Wild Turkey nesting sites and on 5 of the artificial nests that we had placed at random
locations. We baited the artificial nests with 12 unwashed Domestic Chicken
eggs (Hernandez et al. 1997b, Yahner and Mahan 1996). Using binding wire, we
positioned the time-lapse/motion-detecting cameras (Reconyx™ RM45) on tree
trunks within 5 m of the next bowls, and then programmed them to be triggered
once every 5 minutes or by movement near the artificial nest bowl. Based on field
evidence from the cameras, we calculated the percentage of nest-predation events
that involved total predations and partial predations relative to species of predator
and relative to type of predator (mesopredator/other type of predator).
Analyses
We used logistic regression including all variables (year, season, study site, nest
site, and presence of a decoy) to investigate the factors that influenced the likelihood
of predation on artificial nests (Vander Haegen and Degraaf 1996, Vander
Haegen et al. 2002). We also used logistic regression including all variables (year,
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season, study site, nest site, the presence of a decoy, and time to predation) to
investigate relationships between the likelihood of artificial Wild Turkey nests being
preyed on by mesopredators rather than by other predators. We used likelihood
ratio and Hosmer-Lemeshow tests to determine the suitability of the models and
odds ratios (OR) to identify the influence of main effects (Vander Haegen and Degraaf
1996, Yahner and Wright 1985). We used OR to reflect the influence of each
of the significant variables on the likelihood of artificial nest predation.
We recorded the time to predation (from the time of nest deployment) for each
artificial nest. We used fixed-effects analysis of variance (ANOVA; Zar 1999) to
determine whether the time to predation of artificial nests varied relative to type
of predator, year, season, study site, location on historic nest site, and presence of
Wild Turkey decoy.
Results
We monitored 20 artificial Eastern Wild Turkey nests (artificial nests) seasonally
from spring 2009 to spring 2010, and 19 artificial nests from summer 2010
to summer 2011. Confirmed species that depredated artificial Wild turkey nests
were: Corvus brachyrhynchos Brehm (American Crow), Procyon lotor L. (Raccoon),
Didelphis virginiana Kerr (Virginia Opossum), Urocyon cinereoargenteus
Schreber (Gray Fox), Dasypus novemcinctus L. (Nine-banded Armadillo), Lynx
rufus Schreber (Bobcat), Canis latrans Say (Coyote), Picoides sp. (woodpeckers),
and unidentified snakes as well as other unknown nest predators (Table 1). For
mesopredators, the mean times to predation on artificial Wild Turkey nests were
6.5 days (n = 41, SE = 0.72) for Raccoons, and 9.0 days (n = 10, SE = 1.65) for
Opossums; American Crows discovered and preyed on artificial Wild Turkey nests
more quickly (mean = 4.3 days, n = 58, SE = 0.56).
Approximately 61% (118 of 194) of all artificial nests monitored (regardless of
season) were depredated. On study site 1, 53% of artificial nests were depredated,
Table 1. Predators responsible for preying on artificial Wild Turkey nests, in the Pineywoods of East
Texas from spring 2009 to fall 2011.
Seasonal predation
Entire study Winter Spring Summer Fall
Predator n % n % n % n % n %
Armadillo 1 0.8 0 0.0 1 3.0 0 0.0 0 0.0
Bobcat 1 0.8 1 4.8 0 0.0 0 0.0 0 0.0
Coyote 1 0.8 0 0.0 0 0.0 1 2.1 0 0.0
Crow 57 48.3 16 76.2 18 54.5 16 33.3 7 43.8
Gray Fox 2 1.7 0 0.0 2 6.1 0 0.0 0 0.0
Oppossum 10 8.5 1 4.8 0 0.0 6 12.5 3 18.8
Raccoon 41 34.7 2 9.5 11 33.3 22 45.8 6 37.5
Snake 1 0.8 0 0.0 1 3.0 0 0.0 0 0.0
Woodpecker 1 0.8 0 0.0 0 0.0 1 2.1 0 0.0
Unknown 3 2.5 1 4.8 0 0.0 2 4.2 0 0.0
Total 118 21 33 48 16
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compared to 69% on study site 2. In 2011, 85% of all artificial nests were depredated,
compared with 56.7% in 2009 and 62.5% in 2010. 67% of artificial nests deployed
on historic Wild Turkey nest sites were depredated compared, to 54% of randomly
located artificial nests. We found that 86.3% of artificial nests were depredated in
summer, compared with 55% in fall, 54.3% in spring, and 69% in winter.
Study site (study site 1: Z = -2.69, df = 1, P = 0.007), year (2011: Z = 4.19, df = 1,
P < 0.001), nest site (Z = 2.09, df = 1, P = 0.036), and season (summer: Z = 3.30,
df = 1, P < 0.001) influenced whether artificial nests were depredated. Using the
Hosmer-Lemeshow goodness-of-fit test, we failed to detect a significant difference
between the full model and the null model (χ2 = 7.95, df = 8, P = 0.44), indicating
that the model fit the data. We found further support for the plausibility of the model
using a likelihood ratio test (log likelihood = -02.94, df = 9, P < 0.001). The OR
indicated that study site 1, the year 2011, real nest sites, and summer season were
variables that increased the likelihood of artificial nests bein g depredated.
Between spring 2009 and fall 2011, 118 artificial nests were depredated,
53.4% (n = 63) by species other than mesopredators (Table 2). The variables year
(2010: Z = -3.95, df = 1, P < 0.001), year (2011: Z = -2.76, df = 1, P = 0.006),
nest site (Z = 2.31, df = 1, P = 0.02), and time to predation (Z = 3.20, df = 1, P =
0.001) influenced whether an artificial nest was depredated by a mesopredator
or another type of predator. Using a Hosmer-Lemeshow test, there was no difference
between the full model and the null model (χ2 = 9.35, df = 8, P = 0.31), which
indicated that the model fit the data. We found further evidence for the fit of the
model using a likelihood ratio test (log likelihood = -57.44, df = 9, P < 0.001).
The mean time between deployment of an artificial nest and a mesopredator preying
on the nest was 6.8 days (n = 57, SE = 0.63 days), whereas the mean time for
other predators was 4.6 days (n = 60, SE = 0.56 days), mostly because of the relatively
prompt depredation of nests by American Crows.
The mean time to predation in 2009 was 6.1 days (n = 29, SE = 0.92), in 2010 it
was 7.0 days (n = 41, SE = 0.75), and in 2011 it was 4.2 days (n = 47, SE = 0.57).
The relationship between time to predation and year (F = 8.14, df = 2, P < 0.001)
indicated that differences existed in time to predation between 2009 and 2010
Table 2. Contributions of mesopredators to predations on artificial artificial Wild Turkey nests in the
Pineywoods of East Texas from spring 2009 to fall 2011.
Predation
Type of predator Time period n %
Mesopredator 2009–2011 55 46.6
Other 2009–2011 63 53.4
Mesopredator 2009 24 82.8
Other 2009 5 17.2
Mesopredator 2010 13 31.7
Other 2010 28 68.3
Mesopredator 2011 18 37.5
Other 2011 30 62.5
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(Tukey HSD difference = 3.71, P = 0.002), and between 2010 and 2011(Tukey HSD
= -2.92, P = 0.006). Time to predation for mesopredators varied from other nest
predators (F = 6.4, df = 1, P = 0.013). The mean time to predation varied between
study sites (F = 7.91. df = 1, P = 0.006; study site 1: 7.0 days, n = 53, SE = 0.66;
study site 2: 4.5 days, n = 64, SE = 0.52). The variables season (F = 0.141, df = 3,
P = 0.93), nest site (F = 0.203, df = 1, P = 0.65), and the presence of a decoy (F =
2.58, df = 1, P = 0.11) had no influence on time to predation.
Artificial nests were completely depredated during the first visit by a nest predator
in 31% of cases, there was secondary predation in 68% of cases, and in one
instance five of the eggs were eaten and the rest were still in place and intact when
we removed the artificial nest. Only 5% of secondary predations were performed by
species other than those that had initially preyed upon the artificial nests. In three
instances, mesopredators (Raccoon = 2, Gray Fox = 1) were the secondary predator
on artificial nests that Crows had initially depredated, and in one instance an American
Crow was secondary predator at an artificial nest depredated by a snake. There
was little delay between primary and secondary predations (mean = 0.96 days, n =
80, SE = 0.19). The mean times to secondary predation were 0.84 days (n = 28, SE
= 0.25) for Raccoons, 0.68 days (n = 10, SE = 0.1) for Opossums, and 0.59 days (n
= 33, SE = 0.16) for American Crows. In 76% (n = 72) of predations on artificial
nests by mesopredators, there were instances of secondary predation; 60% (n = 38)
of primary predation events by other predators were followed by secondary predation.
On 70% (n = 28) of the occasions that Raccoons preyed on artificial nests,
nests were not completely depredated in the first predation bout, while 63% (n =
36) of artificial nest predations by Crows were followed by seco ndary predations.
Discussion
The degree to which American Crows preyed on artificial nests was greater in
our study within the Pineywoods than in other studies (Baker 1978, Davis 1959,
Hernandez et al. 1997a, Miller and Leopold 1992, Pharris and Goetz 1980). From
January 2009–September 2011, the incidence of predation on artificial nests by
American Crows increased. This finding might be attributable to natural variability
in the degree to which various nest predators prey on nests (Bayne et al. 1997, Buler
and Hamilton 2000, Mezquida 2003). Although we changed the location of artificial
nest sites on an annual basis, we used the same study sites each season throughout the
year, which may have provided sufficient exposure for resident American Crows to
cue in on our placement of the artificial nests (Wilson et al. 1998).
Corvids, such as American Crows, primarily use visual cues to locate nests (Santisteban
et al. 2002). The presence of Wild Turkey hens on live nests camouflages
eggs from avian predators. Despite this, corvids have been recorded preying on
active Wild Turkey nests (Dreibelbis et al. 2008). Wild Turkeys cover their newly
laid eggs prior to the onset of incubation, but once incubation commences, hens do
not camouflage the eggs when they leave the nest for short periods to forage (Healy
1992). We made no attempt to obscure the eggs within the nest bowls, which may
have made the eggs more obvious targets for visual predators. The high visibility
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of the eggs within the nest bowls and the concomitant high incidence of American
Crows depredating artificial nests might be an important factor that differentiates
between predators of artificial Wild Turkey nests and those of real nests, especially
because Wild Turkeys have been observed defending their nests from crows (B.
Collier, Institute for Renewable Natural Resources, Texas A&M University, College
Station, TX, pers. comm.). The relatively short time that it took American
Crows to detect artificial nests is probably also a consequence of the easily visible
eggs in artificial nests when no female Turkey was present.
The diversity of mammalian species that preyed on artificial Wild Turkey nests
was similar to that reported in other US studies (Miller and Leopold 1992), and
more specifically those conducted in the Southeast (Lovell et al. 1995). Raccoons
and Opossums were the most frequent mammalian mesopredators on artificial
Wild Turkey nests in our study. Although we recorded Bobcats and Coyotes at
artificial nests, the instances of predation on artificial nests by these mesopredators
was limited to one incident each. Many studies have reported that mammalian
mesopredators kill adult Wild Turkeys, especially incubating hens (Glidden 1975,
Miller and Leopold 1992, Speake 1980). Yet, we found no evidence of Wild Turkey
feathers, bones, or other remains in Bobcat, Coyote or Raccoon scats found on
these study sites during a concurrent study (Melville 2012). However, there was an
increase in the incidence of predation, specifically by mesopredators, on our study
nests in summer, a pattern that has been attributed to decreases in availability of
small mammalian prey (Melville 2012), and the consequent necessity to increase
the search rate for preferred prey by nest predators during summer (Hoi and Winkler
1994, Knick 1990, Melville 2012, Schmidt 2008).
Wild Turkeys select nest sites that allow them to conceal themselves from predators
(Holbrook et al. 1985, 1987; Isabelle 2010; Lazarus and Porter 1985; Martin
1993b; Porter 1992; Schmutz et al. 1989). We found that during spring, the rate
of predation on historic nest sites was similar to that for all artificial nest sites.
However, when we assessed the degree of predation over all seasons, the level to
which historic nest sites were depredated was greater than for the rates on randomly
located sites. This finding was probably due to the phenology of the plants in the
region: a dense, shrubby understory develops during spring (Chenault 1940, Halls
1973). The dense understory served as cover for the artificial nests, and as the seasons
proceeded, herbaceous plants senesced and provided less nesting cover than
during spring.
We hypothesized that the presence of a Wild Turkey decoy as a proxy for the
presence of a Wild Turkey hen would reduce the incidence of predation on artificial
nests because it increased the level of camouflage for the eggs to protect them from
both avian and terrestrial predators. In addition, the physical presence of the decoy
was unfamiliar to predators and might have caused them to be more cautious when
approaching the artificial nests. This was not the case. Thus, differences in predation
rates between artificial nests and real nests appears to be the consequence of a
combination of cues and not merely the physical presence of a proxy for a hen at
the nest site.
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It was clear from our study that predation on artificial nests was extensive on
both study sites, and our results indicated that nest predation may contribute to
reduced productivity of Eastern Wild Turkeys in the Pineywoods of East Texas.
It is necessary to monitor live Wild Turkey nests to confirm which predators are
responsible for preying on them. To this end, we recommend an ongoing program
to monitor female Eastern Wild Turkeys via radio transmitters in conjunction with
future translocations to help researchers to locate nests and monitor them with nest
cameras. If no future translocations are planned, attempts should be made to capture
extant female Wild Turkeys, fit them with radio transmitters, and monitor their
nests to determine which predator species are responsible for preying on the nests.
Our data could then be compared with the findings from nest-monitoring studies to
determine if our results mirror real-nest predation events.
Acknowledgments
This project was funded by federal excise taxes on sport hunting arms and ammunition,
Grant W132R, with the Texas Parks and Wildlife Department. Support for this research was
also provided by Texas A&M University (Department of Wildlife and Fisheries Science),
and Stephen F. Austin State University (The Arthur Temple College of Forestry and Agriculture).
We thank all the researchers and research technicians including A. Wadyko, J. van
Woert, T. Yurick, J. Deatherage, J.Rogers, J. Isabelle, A. Davis, S. Seidel, and J. Wisnant.
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