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                  <text>The research in this publication was partially or fully funded by Colorado Parks and Wildlife.

Dan Prenzlow, Director, Colorado Parks and Wildlife • Parks and Wildlife Commission: Marvin McDaniel, Chair • Carrie Besnette Hauser, Vice-Chair
Marie Haskett, Secretary • Taishya Adams • Betsy Blecha • Charles Garcia • Dallas May • Duke Phillips, IV • Luke B. Schafer • James Jay Tutchton • Eden Vardy

�Wildlife Society Bulletin 42(4):616–621; 2018; DOI: 10.1002/wsb.935

Original Article

Using Maternal Mule Deer Movements to
Estimate Timing of Parturition and Assist
Fawn Captures
MARK E. PETERSON,1,2 Department of Fish, Wildlife, and Conservation Biology, Colorado State University, 1474 Campus Delivery, Fort Collins,
CO 80523, USA
CHARLES R. ANDERSON, JR, Mammals Research Section, Colorado Parks and Wildlife, 317 W Prospect Road, Fort Collins, CO 80526, USA
MATHEW W. ALLDREDGE, Mammals Research Section, Colorado Parks and Wildlife, 317 W Prospect Road, Fort Collins, CO 80526, USA
PAUL F. DOHERTY, JR, Department of Fish, Wildlife, and Conservation Biology, Colorado State University, 1474 Campus Delivery, Fort Collins,
CO 80523, USA

ABSTRACT Movement patterns of maternal ungulates have been used to determine parturition dates and
aid in locating fawns, which may be important for understanding reproductive rates (e.g., pregnancy and
fetal), but such methods have not been validated for mule deer (Odocoileus hemionus). We first determined
timing of parturition using vaginal implant transmitters (VITs) and then predicted timing of parturition
using VITs in conjunction with Global Positioning System collar data in the Piceance Basin of northwestern
Colorado, USA, during 2012–2014. We examined daily movement rate to determine differences in
movement rate among days (7 days pre- and postpartum) and for movement patterns indicative of
parturition. Mean daily movement rate (m/day) of 102 maternal deer decreased by 46% from 1 day
preparturition (�x ¼ 1,253, SD ¼ 1,091) to parturition date (�x ¼ 682, SD ¼ 574), and remained at this low rate
1–7 days postpartum. We applied an independent data set to validate predicted parturition dates based on
daily movement rate. We estimated day of parturition correctly (i.e., day 0), within 1–3 days postparturition,
and �4 days postparturition of field-reported dates for 10 (29%), 21 (60%), and 4 (11%) maternal females,
respectively. For novel data sets, we predict that a mule deer female whose daily movement rate decreases by
�46% and remains low �3 days postparturition particularly when preceded by a sudden increase in
movement—has given birth. However, we caution that disturbance of deer by field crews should be
minimized, and if birth sites are not found, neonatal mortality will be underestimated. Our results can help
determine timing and general location of parturition as an aid in capturing fawns when the use of VITs is not
feasible, with the ultimate objective of estimating pregnancy, fetal, and fawn survival rates if birth sites are
found. Ó 2018 The Wildlife Society.
KEY WORDS birth, Colorado, Global Positioning System, movement, mule deer, Odocoileus hemionus, vaginal
implant transmitter.

Determining parturition dates and locating birth sites are
important for estimating pregnancy, fetal, and fawn survival rates
(Peterson et al. 2017, 2018), which are needed to accurately
quantify fawn recruitment and comprehend population dynamics of ungulates (Gaillard et al. 1998, Eberhardt 2002, Bonenfant
et al. 2005, Forrester and Wittmer 2013). Maternal deer
movement patterns from Global Positioning System (GPS)
collar data (Long et al. 2009), daily triangulation, and daily
radiolocations from an aircraft have been used to approximate
parturition date and aid in locating fawns (Huegel et al. 1985,
Received: 19 January 2018; Accepted: 14 October 2018
Published: 23 December 2018
1

E-mail: mark.peterson313@gmail.com
Present address: South Dakota Department of Game, Fish and Parks,
4130 Adventure Trail, Rapid City, SD 57702, USA.
2

616

Kunkel and Mech 1994, Carstensen et al. 2003). However,
problems potentially exist for effective location of birth sites,
neonates, and fawns with these methods. First and most
importantly, fawns may not always be located in a timely manner
and stillbirths and early mortalities may not be detected, which
bias survival estimates (Huegel et al. 1985, Kunkel and Mech
1994, Gilbert et al. 2014, Chitwood et al. 2016). Second, when
twins occur, they are not often together and often are missed
when relying upon dam behavior (Carstensen et al. 2003).
Third, use of triangulation and radiolocations requires daily
monitoring that demands extensive logistical and financial
considerations. Last, unnecessary disturbance of maternal–fawn
interactions occur if fawns are not located on the first attempt
and subsequent trips are needed.
To better understand how maternal movement rates reflect
parturition dates, knowing movement and parturition dates
Wildlife Society Bulletin

�

42(4)

�with minimal error is needed. The use of vaginal implant
transmitters (VITs) allow for limited error in determining
parturition dates and locations, but is costly (Bishop et al.
2009, 2011; Carstensen et al. 2009). Relying upon maternal
deer movement rates to identify parturition dates is cheaper
and easier, but has not been validated with VITs. Thus, we
compared GPS collar movement and VIT data to evaluate
the efficacy of determining exact parturition date for mule
deer (Odocoileus hemionus) similar to what has been done for
elk (Cervus elaphus; Vore and Schmidt 2001).
Using movement patterns of maternal ungulates to
determine parturition date is possible because movements
often change substantially after parturition (Huegel et al.
1985, Vore and Schmidt 2001, Long et al. 2009).
Specifically, deer restricted movement rates by approximately
50% to stay within an area 1–7 days after parturition (Huegel
et al. 1985, Long et al. 2009). Based on previous studies
(Huegel et al. 1985, Long et al. 2009), we predicted that
mule deer movement rates would significantly decrease
immediately after parturition and continue to be reduced for
up to 7 days after parturition. Our goal was to apply VITs to
signal birth events and locations and GPS movement data to
assess reliability of GPS data alone to detect mule deer birth
events and aid in capturing fawns. These data can be applied
to facilitate fawn capture efforts and pregnancy and fawn
survival rate estimates without the increased expense and
effort associated with traditional VIT studies if birth sites are
found.

STUDY AREA
We examined daily movement rates (m/day) of maternal
mule deer relative to parturition date in the Piceance Basin of
northwestern Colorado, USA, during 2012–2014. Deer in
this area migrated from low-elevation winter ranges to highelevation summer ranges where they gave birth (Lendrum
et al. 2013). Summer ranges covered parts of Garfield,
Moffat, Rio Blanco, and Routt counties in northwestern
Colorado (39.5808N, �107.9618W and 40.3308N,
�107.0288W) and elevations ranged from 1,900 m to
3,150 m above sea level. Summer range habitat was
dominated by Gambel oak (Quercus gambelii), mountain
mahogany (Cercocarpus montanus), Utah serviceberry (Amelanchier utahensis), mountain snowberry (Symphoricarpos
oreophilus), chokecherry (Prunus virginiana), quaking aspen
(Populus tremuloides), big sagebrush (Artemisia tridentata),
two-needle pinyon pine (Pinus edulis), and Utah juniper
(Juniperus osteosperma). Dominant habitat was interspersed
with Douglas-fir (Pseudotsuga menziesii), Engelmann spruce
(Picea engelmannii), and subalpine fir (Abies lasiocarpa) forests
(Garrott et al. 1987). Shrubs, forbs, and grasses common to
the area have been described (Bartmann 1983, Bartmann
et al. 1992).

METHODS
Adult Female Capture and Handling
In early December 2011–2013, a helicopter crew captured
adult female mule deer (�1.5 yr old) using helicopter netPeterson et al.

�

Estimating Parturition Date of Deer

gunning techniques (Webb et al. 2008, Jacques et al. 2009).
The helicopter crew blindfolded and hobbled deer, and
chemically sedated them with 0.5 mg/kg of Midazolam and
0.25 mg/kg of Azaperone intramuscularly. We fit each
captured deer with a GPS radiocollar (Model G2110D;
Advanced Telemetry Systems, Inc., Isanti, MN, USA)
programmed to obtain locations at 5-hr intervals.
In early March 2012–2014, a helicopter crew recaptured
radiocollared adult females as described for December
captures. For each captured deer, we determined pregnancy
status and number of in utero fetuses using ultrasonography
(SonoVet 2000; Universal Medical Systems, Inc., Bedford
Hills, NY, USA; Stephenson et al. 1995, Bishop et al. 2007).
If an adult female was pregnant, we inserted a VIT (Model
M3930; Advanced Telemetry Systems, Inc.; Bishop et al.
2011, Peterson 2016, Peterson et al. 2018). All capture,
handling, radiocollaring, and VIT insertion procedures were
approved by the Institutional Animal Care and Use
Committee at Colorado Parks and Wildlife (protocol
#17-2008 and #01-2012) and followed guidelines of the
American Society of Mammalogists (Sikes et al. 2016).
Adult Female Monitoring and Location of Birth Sites
During the parturition period (late May–mid-Jul), we
checked radiocollar and VIT signals daily from a Cessna
182 or 185 (Cessna Aircraft Co., Wichita, KS, USA) fixedwing aircraft, weather permitting. Ground crews simultaneously located the radiocollared adult female and VIT by
homing on the ground after we detected a fast pulse rate (80
beats/min) signifying expulsion and presumably birth.
Ground crews then grid searched for fawns and a birth
site near (�400 m) the female and expelled VIT for up to 1 hr
because fawns can leave birth sites within 6 hr of birth
(Johnstone-Yellin et al. 2006, Peterson et al. 2018). Crews
retrieved VITs and recorded location of birth sites using a
hand-held GPS (Garmin GPSMAP 62S, Oregon 650, or
Montana 650; Garmin International Inc., Olathe, KS,
USA).
Daily Movements of Maternal Females and Parturition
Date
We programmed radiocollars deployed on adult females to
release 16 months following initial capture. We retrieved
radiocollars associated with collar drop or mortalities and
downloaded GPS data. We imported data into ArcMap 10.2
(Environmental Systems Research Institute, Inc., Redlands,
CA, USA) to determine locations for each female. To
calculate daily movement when locations overlapped
midnight, we divided the distance moved by 5 and multiplied
it by the number of hours in each day. We analyzed daily
movement rate of maternal females relative to parturition
date and determined exact parturition dates based on VIT
expulsion date. We fixed the parturition date of each female
equal to zero and calculated dates before and after as negative
and positive, respectively. We then calculated daily
movement rate from 7 days pre- and postpartum and
percent change relative to the previous day. Field crews
disturbed females when attempting to confirm birth and
capture fawns, contributing to increased movement after
617

�parturition; thus, we censored one location encompassing the
time of a search for all deer (Peterson et al. 2018).
Methods and Validation Procedure
We examined daily movement rate of each female from
7 days pre- and postpartum to determine differences in
movement rates. Specifically, we looked for patterns
indicative of parturition, including a long-distance movement followed by a significant decrease immediately after
parturition (Huegel et al. 1985, Vore and Schmidt 2001,
Long et al. 2009). We used overlap of 95% confidence
intervals to assess significant differences in movement rates.
To determine the utility of our observations, we used an
independent data set that was collected for an ongoing study
focused on deer demographics in the Piceance Basin
(Anderson 2017) using the same methodology described
above. We used the independent data set from deer captured
in 2015 to estimate day of parturition based on daily
movement rate. We plotted daily movement rate of each deer
from 7 days preparturition to 7 days postparturition. We then
examined these data without knowledge of field-reported
parturition dates and estimated day of parturition using 2
different rules. We estimated day of parturition when daily
movement rate of each deer decreased by �46% and
remained at this low rate for �3 days, particularly when
preceded by a sudden increase in movement (rule 1).
Alternatively, we estimated day of parturition when deer
moved �700 m/day and remained at this low rate for �3 days
(rule 2). Lastly, we evaluated how our estimated parturition
dates compared with field-reported parturition dates.

RESULTS
During March 2012–2014, we documented pregnancy status
of 348 females and inserted VITs in 331 pregnant females.
We were unable to monitor VITs in 124 females for various
reasons (e.g., denied access to private property, VITs
malfunctioned, mortalities before parturition). We located
140 birth sites and found VITs at 120 of those sites.
Movement data were not available for 25 females because of
various reasons (e.g., GPS collar malfunctions, mortalities
before parturition) and we excluded these females from the
analysis as well as 13 females who produced stillborns.
Ultimately, we analyzed movement data for 102 maternal
females (i.e., produced live fawn) from 2012 to 2014.
Daily Movements of Maternal Females
Maternal mule deer moved significantly more 1 day
preparturition (�x ¼ 1,253 m, SD ¼ 1,091 m) than the day
of parturition (�x ¼ 682 m, SD ¼ 574 m; Fig. 1A), a 46%
reduction in mean daily movement rate (Fig. 1B). On
average, maternal deer moved �700 m/day from 1 to 7 days
postparturition (Fig. 1A), but movement varied 1–7 days
preparturition (Fig. 2A), on day of parturition (Fig. 2B), and
1–7 days postparturition (Fig. 2C). Of the maternal females
(n ¼ 102) comprising the 2012–2014 data set, 39 (38%)
made a long-distance movement 1 day preparturition
(�x ¼ 2,344 m, range ¼ 1,336–5,867 m; Fig. 3A) and 36
(35%) made a long-distance movement 2–4 days preparturition (�x ¼ 2,466 m, range ¼ 1,302–10,297 m).
618

Figure 1. Mean daily movement (A; error bars ¼ 95% CI) and change in
movement rate (B) of maternal mule deer from 7 days pre- and
postparturition in the Piceance Basin, Colorado, USA, 2012–2014.

Of the maternal females (n ¼ 35) comprising the independent data set, 18 (51%) made a long-distance movement
1 day preparturition (�x ¼ 5,699 m, range ¼ 1,332–16,356 m;
Fig. 3A) and 11 (31%) made a long-distance movement 2–4
days preparturition (�x ¼ 2,066 m, range ¼ 1,375–3,217 m.
Using rule 1, we estimated day of parturition correctly (i.e.,
day 0), 1–3 days postparturition, and �4 days postparturition
of field-reported dates for 10, 21, and 4 maternal females,
respectively (Table 1). Using rule 2, we estimated day of
parturition correctly, 1–3 days postparturition, and �4 days
postparturition of field-reported dates for 9, 18, and 8
maternal females, respectively (Table 1). Post hoc examination of data showed daily movement rate declined during day
of parturition, but suddenly increased on subsequent days for
females who produced stillborns (Fig. 3B). Barren females
did not exhibit a sustained decline in movement of �46% or
&lt;700 m/day (Fig. 3C).

DISCUSSION
As predicted, maternal mule deer exhibited distinct daily
movement patterns before versus after parturition. Parturient
female deer tend to seek isolation and display restlessness 1–2
days before parturition (Clutton-Brock and Guinness 1975,
Townsend and Bailey 1975), which can contribute to
increased movement just prior to parturition. Thus, we
suggest that a maternal mule deer whose daily movement rate
suddenly increases may be within 1–4 day(s) of giving birth
(i.e., rule 1). We also suggest that a maternal deer whose daily
movement rate decreases by �46% has given birth (i.e., rule
Wildlife Society Bulletin

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42(4)

�Figure 2. Distribution of movement rates of maternal mule deer from 1 to
7 days preparturition (A), on day of parturition (B), and from 1 to 7 days
postparturition (C) in the Piceance Basin, Colorado, USA, 2012–2014.

1). Restricted movement of females after parturition may be
attributable to fawns that are entirely dependent on a hiding
strategy for survival because of vulnerability to predation
(Lent 1974, Geist 1981, Bishop et al. 2009, Monteith et al.
2014, Shallow et al. 2015).
Application of GPS movement data can provide information about mule deer parturition to enhance data collection
efforts addressing pregnancy rates and fawn monitoring. We
suggest that GPS collar data surrounding a parturition
period be plotted to detect a sudden increase and then
sustained reduction in daily movement rate to identify
parturition date. Increased movement prior to parturition is

Figure 3. Examples of mean daily movement rate of mule deer from 7 days
pre- and postparturition representing viable mule deer fawn (A) and stillborn
neonates (B), and a barren female (C) in the Piceance Basin, Colorado, USA,
2012–2014. A sudden and sustained reduction in daily movement rate of
�46% suggests a female has given birth, particularly when preceded by a
sudden increase in movement.

fairly consistent among parturient females and subsequent
births following this behavior are highly likely. Eliminating
or reducing disturbance of deer by field crews is advised to
provide accurate estimates of parturition dates. The recent
advent of 2-way satellite communication and remotely
downloadable collars paired with our method could reduce
aerial and field monitoring and allow for verification of
parturition to estimate pregnancy rate and capture of fawns
without VITs. Other studies have successfully used real-time
movement patterns of radiocollared ungulates without using

Table 1. Differences in estimating day of parturition correctly (i.e., day 0), parturition within 1–3 days postparturition, and �4 days postparturition of fieldreported dates using 2 different rules and likelihood of detecting mule deer fawns within 3 days postparturition, Piceance Basin, Colorado, USA, 2012–2014.
Rulea Day of parturition (%)
1
2
a

29
26

1–3 days postparturition
(%)

�4 days postparturition
(%)

Successful births detected �3 days postparturition
(%)

60
51

11
23

89
77

Rule 1: mean daily movement rate decreased by �46% for 3–4 days, particularly when preceded by a sudden increase in movement. Rule 2: mean daily
movement rate decreased �700 m/day for 3–4 days.

Peterson et al.

�

Estimating Parturition Date of Deer

619

�VITs to document parturition (DeMars et al. 2013, McGraw
et al. 2014) and aid in capture of moose (Alces americanus)
calves (Severud et al. 2015); whereas, we validated movement
patterns of radiocollared mule deer and developed a method
to determine parturition dates and detect birth events to aid
in capture of fawns.
If fetal and fawn survival rates are desired in addition to
parturition date, then a combination of our method with
real-time GPS data could be used to send field-capture crews
in at the correct time to find birth sites (i.e., when daily
movement rate decreases �46%, particularly when preceded
by a long-distance movement). Specifically, a suspected
parturition event could be investigated by locating the
radiocollared female and observing her behavior to determine
whether parturition occurred and aid in capturing fawns and
possibly detecting stillborn neonates and early mortalities
(Huegel et al. 1985, Carstensen et al. 2003). However, use of
GPS data to locate exact birth sites could be difficult without
VITs, so finding stillborns could be limited. Still birth rates
could be underestimated using the GPS data approach and
requires further evaluation. Our approach following rule 1
provided 89% detection of fawn births within 3 days, when
fawn movements are still limited and could provide for
capture and collaring efforts (Jackson et al. 1972, Geist
1981). Capturing fawns within 3 days after birth increases
capture success compared with �4 days after birth. Detection
success could be enhanced by field investigations of reduced
movements immediately following detection from GPS data
(within 1 day) and possibly by using a finer temporal scale
than the 5-hour intervals we used. A finer temporal scale and
real-time monitoring of GPS point clusters may enhance
success in finding birth sites and detecting stillborn neonates
to estimate accurate fetal rates. A study has successfully used
fine-scale movement patterns (i.e., 20-min intervals) of
radiocollared moose without using VITs to identify birth
sites (McGraw et al. 2014). Following our suggestions could
maximize capture of fawns �3 days old to minimize bias in
survival estimates (Gilbert et al. 2014, Chitwood et al. 2016).
Technologically advanced radiocollars and VITs are
increasingly being used to study and comprehend population
dynamics (Hebblewhite and Haydon 2010). Knowledge of
parturition dates and locations and pregnancy and survival
rates are needed to fully comprehend population dynamics of
ungulates (Gaillard et al. 1998, Bonenfant et al. 2005,
Forrester and Wittmer 2013). Ultimately, our proposed
method could help validate such estimates when the use of
VITs is not cost-effective or logistically feasible.

MANAGEMENT IMPLICATIONS
For field biologists and managers, this study provides a
method for determining parturition date and pregnancy rate
for mule deer and potentially other ungulates. Our method
paired with real-time GPS data can also be applied to
facilitate fawn capture and fetal and fawn survival rate
estimates without the increased expense and effort associated
with traditional VIT studies. However, fetal rates can only be
estimated if stillborns are detected and fawns should be
captured close to birth for unbiased estimates of survival. We
620

recommend that managers use the indicator of daily
movement rate decreasing by �46% and remaining low
�3 days postparturition (i.e., rule 1), particularly when
preceded by a sudden increase in movement, to estimate
parturition date and pregnancy and fetal rates. Of importance, our method has the advantages of being simple,
intuitive, and economical, but has the disadvantage of being
qualitative.

ACKNOWLEDGMENTS
We thank L. L. Wolfe, E. J. Bergman, and C. J. Bishop for
assisting with ultrasound and vaginal implant transmitter
insertion. Fixed-wing pilots L. L. Gepfert and L. A. Coulter
provided assistance with aerial telemetry flights and Quicksilver Air, Inc. assisted with deer captures. We thank personnel
from Colorado Parks and Wildlife Area 6, numerous field
technicians, and volunteers for project coordination and
assistance with field work. J. M. Northrup provided assistance
with calculating daily movement rates. E. J. Bergman provided
valuable insight and discussion. M. Festa-Bianchet, S. P.
Haskell, D. B. Johnston, K. A. Logan, P. J. Meiman, G.
Wittemyer, and 1 anonymous referee improved the manuscript
through constructive reviews. Project funding was provided by
ExxonMobil Production/XTO Energy, Williams/WPX Energy, Shell Exploration and Production, EnCana Corporation,
Marathon Oil Corporation, Federal Aid in Wildlife Restoration, the Colorado Mule Deer Foundation, the Colorado Mule
Deer Association, the Boone and Crockett Club, Colorado
Chapter of the Wildlife Society, and the Colorado State
Severance Tax.

LITERATURE CITED
Anderson, C. R., Jr. 2017. Population performance of Piceance basin mule
deer in response to natural gas resource extraction and mitigation efforts to
address human activity and habitat degradation. Federal Aid in Wildlife
Restoration Job Progress Report W-185-R, Colorado Parks and Wildlife,
Fort Collins, USA.
Bartmann, R. M. 1983. Composition and quality of mule deer diets on
pinyon–juniper winter range, Colorado. Journal of Range Management
36:534–541.
Bartmann, R. M., G. C. White, and L. H. Carpenter. 1992. Compensatory
mortality in a Colorado mule deer population. Wildlife Monographs 121.
Bishop, C. J., C. R. Anderson, Jr., D. P. Walsh, E. J. Bergman, P. Kuechle,
and J. Roth. 2011. Effectiveness of a redesigned vaginal implant
transmitter in mule deer. Journal of Wildlife Management 75:1797–1806.
Bishop, C. J., D. J. Freddy, G. C. White, B. E. Watkins, T. R. Stephenson,
and L. L. Wolfe. 2007. Using vaginal implant transmitters to aid in
capture of mule deer neonates. Journal of Wildlife Management
71:945–954.
Bishop, C. J., G. C. White, D. J. Freddy, B. E. Watkins, and T. R.
Stephenson. 2009. Effect of enhanced nutrition on mule deer population
rate of change. Wildlife Monographs 172.
Bonenfant, C., J. M. Gaillard, F. Klein, and J. L. Hamann. 2005. Can we use
the young:female ratio to infer ungulate population dynamics? An
empirical test using red deer Cervus elaphus as a model. Journal of Applied
Ecology 42:361–370.
Carstensen, M., G. D. DelGiudice, and B. A. Sampson. 2003. Using doe
behavior and vaginal-implant transmitters to capture neonate white-tailed
deer in north-central Minnesota. Wildlife Society Bulletin 31:634–641.
Carstensen, M., G. D. DelGiudice, B. A. Sampson, and D. W. Kuehn.
2009. Survival, birth characteristics, and cause-specific mortality of whitetailed deer neonates. Journal of Wildlife Management 73:175–183.
Chitwood, M. C., M. A. Lashley, C. S. DePerno, and C. E. Moorman.
2016. Considerations on neonatal ungulate capture method: potential for
Wildlife Society Bulletin

�

42(4)

�bias in survival estimation and cause-specific mortality. Wildlife Biology:
wlb.00250.
Clutton-Brock, T. H., and F. E. Guinness. 1975. Behaviour of red deer
(Cervus elaphus l.) at calving time. Behaviour 55:287–300.
DeMars, C. A., M. Auger-Methe, U. E. Schlagel, and S. Boutin. 2013.
Inferring parturition and neonate survival from movement patterns of
female ungulates: a case study using woodland caribou. Ecology and
Evolution 3:4149–4160.
Eberhardt, L. L. 2002. A paradigm for population analysis of long-lived
vertebrates. Ecology 83:2841–2854.
Forrester, T. D., and H. U. Wittmer. 2013. A review of the population
dynamics of mule deer and black-tailed deer Odocoileus hemionus in North
America. Mammal Review 43:292–308.
Gaillard, J. M., M. Festa-Bianchet, and N. G. Yoccoz. 1998. Population
dynamics of large herbivores: variable recruitment with constant adult
survival. Trends in Ecology and Evolution 13:58–63.
Garrott, R. A., G. C. White, R. M. Bartmann, L. H. Carpenter, and A. W.
Alldredge. 1987. Movements of female mule deer in northwest Colorado.
Journal of Wildlife Management 51:634–643.
Geist, V. 1981. Behavior: adaptive strategies in mule deer. Pages 157–224 in
O. C. Wallmo, editor. Mule and black-tailed deer of North America.
University of Nebraska Press, Lincoln, USA.
Gilbert, S. L., M. S. Lindberg, K. J. Hundertmark, and D. K. Person. 2014.
Dead before detection: addressing the effects of left truncation on survival
estimation and ecological inference for neonates. Methods in Ecology and
Evolution 5:992–1001.
Hebblewhite, M., and D. T. Haydon. 2010. Distinguishing technology from
biology: a critical review of the use of GPS telemetry data in ecology.
Philosophical Transactions of the Royal Society B: Biological Sciences
365:2303–2312.
Huegel, C. N., R. B. Dahlgren, and H. L. Gladfelter. 1985. Use of doe
behavior to capture white-tailed deer fawns. Wildlife Society Bulletin
13:287–289.
Jackson, R. M., M. White, and F. F. Knowlton. 1972. Activity patterns of
young white-tailed deer fawns in south Texas. Ecology 53:262–270.
Jacques, C. N., J. A. Jenks, C. S. Deperno, J. D. Sievers, T. W. Grovenburg,
T. J. Brinkman, C. C. Swanson, and B. A. Stillings. 2009. Evaluating
ungulate mortality associated with helicopter net-gun captures in the
northern Great Plains. Journal of Wildlife Management 73:1282–1291.
Johnstone-Yellin, T. L., L. A. Shipley, and W. L. Myers. 2006.
Effectiveness of vaginal implant transmitters for locating neonatal mule
deer fawns. Wildlife Society Bulletin 34:338–344.
Kunkel, K. E., and L. D. Mech. 1994. Wolf and bear predation on whitetailed deer fawns in northeastern Minnesota. Canadian Journal of Zoology
72:1557–1565.
Lendrum, P. E., C. R. Anderson, Jr., K. L. Monteith, J. A. Jenks, and R. T.
Bowyer. 2013. Migrating mule deer: effects of anthropogenically altered
landscapes. PLoS ONE 8:e64548.

Peterson et al.

�

Estimating Parturition Date of Deer

Lent, P. C. 1974. Mother–infant relationship in ungulates. Pages 14–55 in
V. Geist and F. R. Walther, editors. The behaviour of ungulates and its
relation to management. International Union for Conservation of Nature
and Natural Resources, Morges, Switzerland.
Long, R. A., J. G. Kie, R. T. Bowyer, and M. A. Hurley. 2009. Resource
selection and movements by female mule deer Odocoileus hemionus: effects
of reproductive stage. Wildlife Biology 15:288–298.
McGraw, A. M., J. Terry, and R. Moean. 2014. Pre-parturition movement
patterns and birth site characteristics of moose. Alces 50:93–103.
Monteith, K. L., V. C. Bleich, T. R. Stephenson, B. M. Pierce, M. M.
Conner, J. G. Kie, and R. T. Bowyer. 2014. Life-history characteristics of
mule deer: effects of nutrition in a variable environment. Wildlife
Monographs 186.
Peterson, M. E. 2016. Reproductive success, habitat selection, and neonatal
mule deer mortality in a natural gas development area. Dissertation,
Colorado State University, Fort Collins, USA.
Peterson, M. E., C. R. Anderson, Jr., J. M. Northrup, and P. F. Doherty, Jr.
2017. Reproductive success of mule deer in a natural gas development area.
Wildlife Biology 2017:wlb.00341.
Peterson, M. E., C. R. Anderson, Jr., J. M. Northrup, and P. F. Doherty, Jr.
2018. Mortality of mule deer fawns in a natural gas development area.
Journal of Wildlife Management 82:1135–1148.
Severud, W. J., G. D. DelGiudice, T. R. Obermoller, T. A. Enright, R. G.
Wright, and J. D. Forester. 2015. Using GPS collars to determine
parturition and cause-specific mortality of moose calves. Wildlife Society
Bulletin 39:616–625.
Shallow, J. R. T., M. A. Hurley, K. L. Monteith, and R. T. Bowyer. 2015.
Cascading effects of habitat on maternal condition and life-history
characteristics of neonatal mule deer. Journal of Mammalogy 96:194–205.
Sikes, R. S., and the Animal Care and Use Committee of the American
Society of Mammalogists. 2016. Guidelines of the American Society of
Mammalogists for the use of wild mammals in research. Journal of
Mammalogy 97:663–688.
Stephenson, T. R., J. W. Testa, G. P. Adams, R. G. Sasser, C. G. Schwartz,
and K. J. Hundertmark. 1995. Diagnosis of pregnancy and twinning in
moose by ultrasonography and serum assay. Alces 31:167–172.
Townsend, T. W., and E. D. Bailey. 1975. Parturitional, early maternal, and
neonatal behavior in penned white-tailed deer. Journal of Mammalogy
56:347–362.
Vore, J. M., and E. M. Schmidt. 2001. Movements of female elk during
calving season in northwest Montana. Wildlife Society Bulletin
29:720–725.
Webb, S. L., J. S. Lewis, D. G. Hewitt, M. W. Hellickson, and F. C. Bryant.
2008. Assessing the helicopter and net gun as a capture technique for
white-tailed deer. Journal of Wildlife Management 72:310–314.

Associate Editor: Haskell.

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              <text>&lt;span&gt;Movement patterns of maternal ungulates have been used to determine parturition dates and aid in locating fawns, which may be important for understanding reproductive rates (e.g., pregnancy and fetal), but such methods have not been validated for mule deer (&lt;/span&gt;&lt;i&gt;Odocoileus hemionus&lt;/i&gt;&lt;span&gt;). We first determined timing of parturition using vaginal implant transmitters (VITs) and then predicted timing of parturition using VITs in conjunction with Global Positioning System collar data in the Piceance Basin of northwestern Colorado, USA, during 2012–2014. We examined daily movement rate to determine differences in movement rate among days (7 days pre- and postpartum) and for movement patterns indicative of parturition. Mean daily movement rate (m/day) of 102 maternal deer decreased by 46% from 1 day preparturition (&lt;/span&gt;&lt;span&gt; = 1,253, SD = 1,091) to parturition date (&lt;/span&gt;&lt;span&gt; = 682, SD = 574), and remained at this low rate 1–7 days postpartum. We applied an independent data set to validate predicted parturition dates based on daily movement rate. We estimated day of parturition correctly (i.e., day 0), within 1–3 days postparturition, and ≥4 days postparturition of field-reported dates for 10 (29%), 21 (60%), and 4 (11%) maternal females, respectively. For novel data sets, we predict that a mule deer female whose daily movement rate decreases by ≥46% and remains low ≥3 days postparturition particularly when preceded by a sudden increase in movement—has given birth. However, we caution that disturbance of deer by field crews should be minimized, and if birth sites are not found, neonatal mortality will be underestimated. Our results can help determine timing and general location of parturition as an aid in capturing fawns when the use of VITs is not feasible, with the ultimate objective of estimating pregnancy, fetal, and fawn survival rates if birth sites are found.&lt;/span&gt;</text>
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              <text>&lt;p&gt;Peterson, M. E., C. R. Anderson Jr, M. W. Alldredge, and P. F. Doherty Jr. 2018. Using maternal mule deer movements to estimate timing of parturition and assist fawn captures. Wildlife Society Bulletin 42:616–621. &lt;a href="https://doi.org/10.1002/wsb.935" target="_blank" rel="noreferrer noopener"&gt;https://doi.org/10.1002/wsb.935&lt;/a&gt;&lt;/p&gt;</text>
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