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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

�Migration Patterns of Adult Female Mule Deer in Response to Energy Development
Charles R. Anderson, Jr.
Colorado Parks and Wildlife
Fort Collins, Colorado
Chad J. Bishop
Colorado Parks and Wildlife
Denver, Colorado
Introduction
Migration is an adaptive strategy that enables animals to enhance resource availability and reduce
risk of predation at a broad geographic scale. Ungulate migrations generally occur along traditional
routes, many of which have been disrupted by anthropogenic disturbances. Spring migration in ungulates
is of particular importance for conservation planning because it is closely coupled with timing of
parturition. The degree to which oil and gas development affects migratory patterns, and whether ungulate
migration is sufficiently prepared to compensate for such changes, has recently been investigated in
Colorado and Wyoming (Lendrum et al. 2012, 2013; Sawyer et al. 2012).
Lendrum et al. (2012, 2013) and Sawyer et al. (2012) address mule deer (Odocoileus hemionus)
migration patterns in relation to energy development from northwest Colorado and south-central
Wyoming, respectively. We address results from the Colorado and Wyoming studies and then compare
similarities and differences. Management and conservation implications are proposed for consideration
and future investigation.
Piceance Basin Mule Deer Migration
Lendrum et al. (2012, 2013) investigated spring migration patterns of adult female mule deer in
the Piceance Basin of northwest Colorado from 2008 to 2010. They used Global Positioning System
(GPS) collars (five location attempts per day) to address habitat use patterns and factors influencing
timing and synchrony of spring migration by comparing areas with ongoing natural gas development
activity to areas with little to no development (Lendrum et al. 2012, n = 167; Lendrum et al. 2013, n =
205). Mean migration distances among study areas varied from 36 to 53 kilometers (distance traveled; n =
4 winter range study areas), averaging 36 kilometers between seasonal ranges (linear distance; study area
range: 3240 km). Piceance Basin mule deer demonstrated rapid spring migration exhibiting median
durations of three to eight days among areas. Stopover use (areas used to increase energy reserves during
migration) along migration paths was rare for Piceance Basin mule deer. Well pad densities along
migration paths within the two developed study areas were 1.5 to 2.0 pads per square kilometer.
Mule deer migrated more quickly through the most developed areas compared with deer in less
developed areas. Additionally, deer migrating through the most developed study areas tended to select
habitat types that provided greater amounts of concealment cover, whereas deer from the least developed
areas tended to select habitats that increased access to forage and cover. Deer selected habitats closer to
well pads and avoided roads in all instances except along the most highly developed migratory routes,
where road densities may have been too high for deer to avoid roads without deviating substantially from
established migration routes.
Environmental factors influencing timing and synchrony of spring migration included snow depth
and emerging vegetation, which varied among years but was highly synchronous among study areas
within years. Migration timing was also influenced by development disturbance, rate of travel, distance
traveled, and late-winter body condition. Rates of travel were more rapid over shorter migration distances
in areas of high natural gas development resulting in delayed departure—but early arrival for females.

Transactions of the 79th North American Wildlife and Natural Resources Conference 
47

�These results indicate that behavioral tendencies to avoid anthropogenic disturbance can be
overridden during migration by the strong fidelity mule deer demonstrate towards migration routes. If
avoidance is feasible, then deer may select areas further from development, whereas in highly developed
areas, deer may simply increase their rate of travel along established migration routes.
Atlantic Rim Mule Deer Migration
Sawyer et al. (2012) used GPS data (location attempts every 2.5 hours) collected from two
subpopulations of mule deer (n = 97) in the Atlantic Rim region of Wyoming to evaluate how different
densities of gas development (coal-bed methane) influenced migratory behavior, including movement
rates and stopover use at the individual level and intensity of use and width of migration route at the
population level. They characterized the functional landscape of migration routes as either stopover
habitat or movement corridors and examined how the observed behavioral changes affected the
functionality of the migration route in terms of stopover use. Atlantic Rim mule deer exhibited relatively
longer migration duration averaging about three weeks, with distances averaging 40 kilometers between
seasonal ranges, and common stopover use along migration paths. Well pad densities were more
concentrated and higher than in the Piceance Basin increasing from 0.8 to 2.8 pads per square kilometer
in the most developed study area.
Sawyer et al. (2012) found migratory behavior to vary with development intensity. They suggest
that mule deer can migrate through moderate levels of development without any noticeable effects on
migratory behavior. However, in areas with more intensive development, animals often detoured from
established routes, increased their rate of movement, and reduced stopover use, while the overall use and
width of migration routes decreased.
In contrast to impermeable barriers that impede animal movement, semipermeable barriers allow
animals to maintain connectivity between their seasonal ranges. Their results identify the mechanisms
(e.g., detouring, increased movement rates, reduced stopover use) by which semipermeable barriers affect
the functionality of ungulate migration routes and emphasize that the management of semipermeable
barriers may play a key role in the conservation of migratory ungulate populations.
Discussion
Environmental conditions were similar between study areas, whereas development intensity and
migratory behavior differed in some respects (Table 1). Migration distances, elevation gradients, and
general habitat types were similar (Table 1), but overstory cover was typically higher in the Piceance
Basin where migratory mule deer took advantage of security cover to avoid development activity, without
detectable deviation from migration paths. Migratory mule deer in both areas traveled more quickly
through developed landscapes, but permeability of migration routes was only inhibited at the more
concentrated development intensity evident in Atlantic Rim, Wyoming. Nonetheless, increased movement
rates through developed areas can discourage use of stopover habitat and reduce the ability of animals to
optimally forage and track vegetation phenology. Whether such behavioral changes have demographic
consequences is unknown, but given the importance of summer nutrition for body condition and
reproduction, any lost foraging opportunities during migration have the potential to incur energetic and
demographic costs and the resulting effect may act as de facto habitat loss. Increased energetic costs
associated with strong deviations in traditional migration routes, and reduced energy intake resulting from
poor timing of arrival on summer range relative to forage conditions, could compromise long-term fitness
of migratory mule deer populations. Thus, conservation measures may be warranted in areas where
expansive and concentrated development activities occur or are planned within the range of long-distance
migratory ungulates.
Interesting differences between the two migratory mule deer populations, which likely was not
related to energy development activities, included the relatively rapid migration duration and reduced
stopover use exhibited by Piceance Basin, Colorado, mule deer (Table 1). The reason for these differences
Special Session Two: Migration Patterns of Adult Female Mule Deer
48 

�is unclear, but could be related to forage conditions and mule deer body condition prior to migration.
Lendrum et al. (2013) noted that mule deer in relatively good condition migrated earlier than deer in poor
condition, which required improved body condition prior to long-distance movements, and it is intuitive
(although speculative) that individuals with improved energy reserves could migrate more quickly
without stopping along the way to “refuel.” It may also be that stopover use in Wyoming reflected an
optimal foraging strategy relative to the timing of green-up as deer progressed in elevation. Where
stopover use is common, identifying and incorporating stopover sites into energy development planning is
critical to sustaining migratory ungulate populations (Sawyer et al. 2012).
Implications
The interactions between migratory mule deer and energy development identified by Lendrum et
al. (2012, 2013) and Sawyer et al. (2012) suggest mule deer may benefit from energy development
planning by considering thresholds of development that may alter migratory behavior. It appears that
migration rate, migration routes, and stopover use, if present, may be altered at high development
intensities. In addition, migratory mule deer may benefit by maintaining security cover along migration
paths, and improved habitat conditions may facilitate more direct and rapid migration requiring less
energy to complete migration. Enhancing permeability along migration routes by applying dispersed
development plans (&lt;2 well pads/km2) and minimizing disturbance to vegetation types by maintaining
security cover should reduce impacts to migratory mule deer as well as other migratory ungulates. Where
feasible, habitat improvement projects on winter range and possibly stopover sites would also enhance
migratory mule deer populations by enhancing energy reserves for long-distance movements and
parturition shortly after summer range arrival. Where possible, directional drilling could be used to extract
energy resources from underneath migration routes while maintaining no surface occupancy. Lastly, we
emphasize that GPS studies now allow managers to accurately map migration routes for entire
populations and identify relatively narrow corridors that are most heavily used thus allowing for the
identification of the most important corridors for migrating ungulates. Where available, we encourage
agencies to incorporate such migration corridors into land-use plans (e.g., resource management plans)
and National Environmental Policy Act documents.
References
Lendrum, P. E., C. R. Anderson, Jr., K. L. Monteith, J. A. Jenks, and R. T. Bowyer. “Migrating Mule
Deer: Effects of Anthropogenically Altered Landscapes.” PLOS ONE 8(5): e64548, 2013.
Lendrum, P. E., C. R. Anderson, Jr., R. A. Long, J. G. Kie, and R. T. Bowyer. “Habitat
Selection by Mule Deer During Migration: Effects of Landscape Structure and Natural Gas
Development.” Ecosphere 3: art82, 2012.
Sawyer, H., M. J. Kauffman, A. D. Middleton, T. A. Morrison, R. M. Nielson, and T. B. Wycoff. “A
Framework for Understanding Semi-Permeable Barrier Effects on Migratory Ungulates.” Journal
of Applied Ecology 50 (2012): 68-78.
Citation: Anderson, C. R., Jr., and C. J. Bishop. 2014. Migration patterns of adult female mule deer in response
to energy development. Pages 47-50 in R. A. Coon &amp; M. C. Dunfee, editors. Transactions of the 79th North
American Wildlife and Natural Resources Conference. Wildlife Management Institute, Gardners, PA, USA.
ISSN 0078-1355.

Transactions of the 79th North American Wildlife and Natural Resources Conference 
49

�Table 1. Comparison of two migratory mule deer populations from Piceance Basin, Colorado
(Lendrum et al. 2012, 2013), and Atlantic Rim, Wyoming (Sawyer et al. 2012), in relation to
environmental conditions, migration behavior, and well pad density of developed landscapes along
migration paths.

Piceance Basin, CO
Atlantic Rim, WY
36 km
40 km
1,980–2,400 m
2,065–2,385 m
PJ woodland, mtn. shrub,
Sparse PJ/sage, sage,
Aspen/conifer
Aspen/sage
Stopover use
Rare
Common
Well pad densitya
1.52.0/km2
0.8–2.8/km2
a
Well pad densities in the Piceance Basin, Colorado, were averaged along entire migration paths of the
two developed study areas (Lendrum et al. 2012). Well pad densities in Atlantic Rim, Wyoming,
represent phased development over a five-year period within a concentrated area along the migration
corridor of the most developed study area (Sawyer et al. 2012).
Mean dist. between seasonal ranges
Range in elevation
General habitat types

50 
Special Session Two: Migration Patterns of Adult Female Mule Deer

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