ArticlePDF Available

Space use and movement patterns of translocated bighorn sheep

Authors:
  • South Dakota Department of Game, Fish, and Parks

Abstract and Figures

Ungulate species have consistently been a major focus of reintroductions to their native ranges. Bighorn sheep (Ovis canadensis) are an ecologically sensitive species, and have experienced population declines throughout their historic range; bighorn sheep inhabited the Black Hills region of South Dakota but were extirpated from the area due to anthropogenic impacts in the early 1900s. To continue to restore populations to the area, we translocated 26 bighorn sheep from Alberta, Canada to the Deadwood Region of the Black Hills. Bighorn sheep were fitted with VHF or GPS collars and monitored throughout the duration of the study (Feb 2015–Jan 2017). Our objectives were to evaluate movement patterns post-release of bighorn sheep in the translocated Deadwood bighorn sheep herd. We utilized 3 types of home-range analyses based on collar data; kernel density estimation (KDE), minimum convex polygon (MCP), and Brownian Bridge Movement Models (BBMM) were used to estimate home-ranges year 1, year 2, and for the duration of the study. Home-range size utilizing KDE (95%; \(\overline{x}\) = 41.41 km2, SE = 10.50), minimum convex polygon (95%; \(\overline{x}\) = 55.73 km2, SE = 15.04), and BBMM (95%; \(\overline{x}\) = 32.95 km2, SE = 4.67) differed among methods. Year 1 home-range sizes (95% BBMM; \(\overline{x}\) = 40.01 km2) were larger than year 2 (95% BBMM; \(\overline{x}\) = 4.08 km2) home-range sizes. Travel distances were also larger in year 1 (\(\overline{x}\) = 431.80 km) than year 2 (\(\overline{x}\) = 368.77 km). Our results indicate that after an acclimation period, which included individual dispersal, the translocated Deadwood bighorn sheep herd settled into smaller home-ranges near the release site.
Content may be subject to copyright.
Vol.:(0123456789)
1 3
Mammalian Biology
https://doi.org/10.1007/s42991-021-00107-4
ORIGINAL ARTICLE
Space use andmovement patterns oftranslocated bighorn sheep
TyJ.Werdel1,2 · JonathanA.Jenks1· JohnT.Kanta3· ChadwickP.Lehman4· TeresaJ.Frink5
Received: 5 June 2020 / Accepted: 25 January 2021
© Deutsche Gesellschaft für Säugetierkunde 2021
Abstract
Ungulate species have consistently been a major focus of reintroductions to their native ranges. Bighorn sheep (Ovis canaden-
sis) are an ecologically sensitive species, and have experienced population declines throughout their historic range; bighorn
sheep inhabited the Black Hills region of South Dakota but were extirpated from the area due to anthropogenic impacts in
the early 1900s. To continue to restore populations to the area, we translocated 26 bighorn sheep from Alberta, Canada to
the Deadwood Region of the Black Hills. Bighorn sheep were fitted with VHF or GPS collars and monitored throughout the
duration of the study (Feb 2015–Jan 2017). Our objectives were to evaluate movement patterns post-release of bighorn sheep
in the translocated Deadwood bighorn sheep herd. We utilized 3 types of home-range analyses based on collar data; kernel
density estimation (KDE), minimum convex polygon (MCP), and Brownian Bridge Movement Models (BBMM) were used
to estimate home-ranges year 1, year 2, and for the duration of the study. Home-range size utilizing KDE (95%;
x
= 41.41
km2, SE = 10.50), minimum convex polygon (95%;
x
= 55.73 km2, SE = 15.04), and BBMM (95%;
x
= 32.95 km2, SE = 4.67)
differed among methods. Year 1 home-range sizes (95% BBMM;
x
= 40.01 km2) were larger than year 2 (95% BBMM;
x
= 4.08 km2) home-range sizes. Travel distances were also larger in year 1 (
x
= 431.80km) than year 2 (
x
= 368.77km). Our
results indicate that after an acclimation period, which included individual dispersal, the translocated Deadwood bighorn
sheep herd settled into smaller home-ranges near the release site.
Keywords Bighorn sheep· Black hills· Home-range· Movement patterns· Resource selection· Translocation· Ovis
canadensis
Introduction
Ungulates have often been the focus of restoration in
response to diminishing populations, due to their economic
value through hunting and ecotourism (Gordon etal. 2004);
between 1973 (Endangered Species Act became law) and
1986, 90% of translocated species were game species and
39% were ungulates (Griffith etal. 1989). Reintroduc-
tions via translocation [intentionally moving individuals
of a species to a different area (Krausman and Cox 2019)]
are executed in localized areas where a species has been
completely extirpated or where low species’ populations
require increased abundance or genetic diversity to remain
viable (Buechner 1960). Evaluating ungulate translocations
is critical, due to expected initial mortality post-release,
relatively low number of individuals initially translocated,
unproven habitat capability, the potential for predation, and
time necessary for acclimation. Results of many reintroduc-
tions and translocations are inadequately detailed, leaving
stakeholders without an understanding of project benefits
Handling editor: Luca Corlatti.
* Ty J. Werdel
werdel@ksu.edu
1 Department ofNatural Resource Management, Edgar S.
Mcfadden Biostress Lab, South Dakota State University,
Brookings, SD57007, USA
2 Present Address: Department ofHorticulture andNatural
Resources, Kansas State University, Manhattan, KS66506,
USA
3 South Dakota Game, Fish andParks, 4130 Adventure Trail,
RapidCity, SD57702, USA
4 South Dakota Game, Fish andParks, 13329 US Highway
16A, Custer, SD57730, USA
5 Department ofApplied Sciences, Burkhiser Complex,
Chadron State College, Chadron, NE69337, USA
T.J.Werdel et al.
1 3
and challenges (Gogan 1990; Zimmerman 2008). Transloca-
tions should be vigorously evaluated to provide information
on carrying capacity, post-release pioneering (i.e., explora-
tory movements), habitat selection, and home range to help
increase translocation success (Douglas and Leslie 1999;
Zimmerman 2008). Pioneering from translocation release
sites allows individuals to acquaint themselves with novel
environments, however, these initial exploratory movements
may lead to increased predation risk (Nicholson etal. 1997;
Scillitani etal. 2013).
Home range has been defined as: “That area traversed
by an individual in its normal activities of food gathering,
mating, and caring for young. Occasional sallies outside the
area, perhaps exploratory in nature, should not be consid-
ered part of the home range (Burt 1943).’’ Burt (1943) did
not explain how to measure or estimate an animal’s home
range, and although this fundamental concept is difficult to
define empirically, ecologists have continually attempted to
accurately analyze movement patterns based on this defini-
tion (White and Garrott 1990; Powell and Mitchell 2012).
With advances in telemetry and global positioning satellite
(GPS) transmitters utilized for marking animal locations,
researchers now commonly estimate home ranges by esti-
mating densities of these locations across the landscape
(Laver and Kelly 2008; Powell 2000; Powell and Mitchell
2012). Traditionally, there have been 3 methods of estimat-
ing home range size for ungulates; kernel density estimator
(KDE) (Worton 1989), minimum convex polygon (MCP)
(Krausman etal. 1989), and Brownian Bridge Movement
Models (BBMM) (Zimmerman 2008; Jacques etal. 2009;
Kie etal. 2010; Wilckens 2014; Parr 2015).
MCP home ranges are becoming a method of the past,
due to the method’s tendency to overestimate true home
ranges (Getz and Wilmers 2004; Kie etal. 2010; Wilckens
2014; Parr 2015) and are known to introduce unpredict-
able biases in small sample sizes by tending to be under-
dispersed (Nilsen etal. 2008). MCP home ranges are now
generally reserved for comparative purposes (between stud-
ies conducted before advances in home range statistical
analyses and contemporary studies) and within single-pop-
ulation studies (Borger etal. 2006). The MCP method also
provides unpredictable changes in variance with increased
sampling effort and temporal variation (Borger etal. 2006).
The most condemnatory aspect of the MCP method is that
it produces strong, likely inadequate, biological assumptions
by determining home range utilizing only data points of the
outermost locations (Worton 1995; Borger etal. 2006). The
KDE method describes the animal’s home range as a utili-
zation distribution by creating multi-dimensional relative
frequency distributions of locations; allowing for estimation
and detection of core areas utilized by the focal population
(Worton 1989; Kie etal. 2010). This is accomplished by
calculating differing densities (i.e., 50%, 95%, 99%) within
the analysis (Barg etal. 2005) and has been shown in simu-
lation studies to represent home range more reliably than
MCP (Worton 1987). KDE methods are utilized due to
their nonparametric nature and ability to produce consistent
results when smoothing and bandwidth selections are simi-
lar (Worton 1989; Kie etal. 2010; Schuler etal. 2014; Parr
2015). However, both smoothing (fixed and adaptive) and
method of bandwidth selection are highly influential in the
final KDE estimates, making comparisons (between studies)
complicated if these parameters are not similar (Laver and
Kelly 2010). Due to the recent prominence of GPS collar
technology in wildlife research, BBMM is increasingly used
because of its ability to delineate true areas of use and define
exploratory movements (Kie etal. 2010; Walter etal. 2011;
Wilckens 2014; Parr 2015). Movement trajectories described
by BBMM may give insight into biologically important
space-use by the focal species (Kie etal. 2010). The BBMM
helps delineate true space-use by filling in space between
sequential locations, irrespective of the density of locations;
the width of the Brownian bridge is conditioned only on the
time duration between the beginning and ending locations
for each pair of locations (Walter etal. 2011). When increas-
ing the density of data points utilized (i.e., 95–99%), BBMM
does not over-smooth the utilization density, but may more
accurately define the area of use for some species that travel
considerable distances (i.e., large ungulates) (Lewis 2007;
Walter etal. 2011).
Bighorn sheep (Ovis canadensis), a North American moun-
tain ungulate, are gregarious, but segregate sexually; males
(hereafter, rams) can occupy habitats with higher predator [i.e.,
coyotes (Canis latrans), mountain lions (Puma concolor)] den-
sities and forage quality, whereas females (hereafter, ewes)
occupy habitats closer to water sources and generally occur
in larger social groups than do rams (Geist 1971; Bleich etal.
1997). Pregnant ewes migrate between two distinct range types
prior to lambing (May–June); moving from low elevation win-
ter ranges with high forage quality, to high elevation lambing
ranges with lower forage quality to avoid predation on new-
born lambs (Festa-Bianchet 1988). Ewes learn home ranges
from maternal bands and may stay with their mothers for up to
6years (Geist 1971; Festa-Bianchet 1988). Rams remain with
their mothers from 1 to 4years before searching out adult ram
bands and forming their own home ranges (Geist 1971; Festa-
Bianchet 1988). Escape terrain, including rocky outcrops and
slopes ≥ 40 degrees (Zimmer man 2008), is a critical habitat
attribute for bighorn sheep (Festa-Bianchet 1986). At Badlands
National Park, Zimmerman (2008) found resident and intro-
duced bighorn sheep were, on average, < 150m from escape
terrain, resulting in linear home ranges that were closely asso-
ciated with badland formations. Northern populations of big-
horn sheep rarely disperse or colonize new areas once home-
ranges are established (Singer etal. 2000a, b; Zimmerman
2008). However, young rams may disperse as a function of
Space use andmovement patterns oftranslocated bighorn sheep
1 3
the pursuit of breeding advantages, while young ewes seldom
disperse due to their tendency to adopt the home-ranges of
adult maternal ewes (Geist 1971; Zimmerman 2008). Disper-
sal by bighorn sheep into unfamiliar or inhospitable habitat
may increase an individual’s susceptibility to predation, stress,
or malnutrition (Van Vuren 1998; Zimmerman 2008). Expe-
ditious establishment of home-ranges near the reintroduc-
tion site, with minimal dispersion from suitable terrain, will
maximize the survivability of translocated bighorn sheep (Van
Vuren 1998; Zimmerman 2008).
Bighorn sheep were once abundant, numbering in the
millions, and occupied habitats across the western United
States, Canada, and Mexico (Buechner 1960). Anthro-
pogenic factors such as uncontrolled harvest, introduced
disease and forage competition from domestic livestock,
reduced and fragmented habitat, and loss of movement cor-
ridors led to major declines in bighorn sheep populations
in the late 19th through the mid-twentieth century (Buech-
ner 1960; Douglas and Leslie 1999; Beecham etal. 2007).
In South Dakota, bighorn sheep historically inhabited the
entire Black Hills region, but the species was extirpated from
the area in the early 1900s (Seton 1929; Zimmerman 2008;
Witte and Gallagher 2012; Parr 2015). The Deadwood area
of the northern Black Hills was vacant of bighorn sheep but
was deemed potentially suitable for reintroduction, based
on habitat suitability models and a qualitative assessment
of topography, forage, and water (SDGFP 2013). However,
despite a prerelease evaluation of the release site, no infor-
mation was available regarding the historical use of the area
by bighorn sheep (SDGFP 2013).
To expand bighorn sheep restoration in the Black Hills,
we translocated 26 bighorn sheep from the Luscar Mine
near Hinton, Alberta, Canada to the Deadwood region of
the northern Black Hills. In an effort to evaluate a critical
large mammal translocation, we investigated space-use and
movement patterns post-release of bighorn sheep in the
translocated Deadwood bighorn sheep herd. Specifically, we
tested the following predictions: (1) translocated bighorn
sheep would engage in a short pioneering period, followed
by the establishment of permanent home-ranges near the
release site, (2) pregnant ewes would utilize separate, rugged
areas, during the lambing period, and (3) BBMM would be
the most accurate home-range estimator for delineating true
space-use of a large ungulate exhibiting pioneering behavior
post-release.
Materials andmethods
Study area
Bighorn sheep were captured on the Luscar Mine in Alberta,
Canada [5,880,593, 473,136, 11N (a reclaimed mine habitat
utilized by a population of ~ 1000 bighorn sheep and the
source of > 350 bighorn sheep translocated throughout
North America) (Teck 2012)]. Indicative of reclaimed
mines, benched high walls have been maintained to pro-
vide escape terrain in proximity to reclaimed grasslands
(MacCallum and Geist 1992). Elevation varied from 1680
to 1860m, including rugged slopes of varying aspect and
angles (Maccallum 2012). Captured bighorn sheep were
then translocated to our study area, the Deadwood region,
located in the northern Black Hills in western South Dakota,
USA (44°2056.55"N, – 103°4228.64"W) (Fig.1). This area
encompasses approximately 8177ha of public (5203ha) and
private (2974ha) land and is located immediately adjacent
to the Deadwood, Lead, and Central City communities in
Lawrence County, South Dakota (SDGFP 2013). Eleva-
tions range from 1073 to 2209m above mean sea level. The
Deadwood region of the Black Hills occurs within the cen-
tral core of the Black Hills, which is typified by canyons,
mountain peaks, and broad valleys (Hoffman and Alexander
1960), and includes areas of reclaimed mining operations
(i.e., Homestake Mine, Gilt Edge Mine). Soils of the region
include limestones, dolomites, and sandstones of Paleo-
zoic origin (Hoffman and Alexander 1960). This region of
the Black Hills receives more precipitation in the form of
snow (156cm) and is cooler (−5°C) than more southern
areas within the Black Hills (Hoffman and Alexander 1960;
NOAA 2017).
Ponderosa pine (Pinus ponderosa) was the dominant
overstory tree species of the region; it occurs in monotypic
stands and was intermixed with small stands of quaking
aspen (Populus tremuloides) and paper birch (Betula papy-
rifera) (Mcintosh 1949; Orr 1959; Hoffman and Alexan-
der 1960; Thilenius 1972; Richardson and Peterson 1974).
Common plant species included Kentucky bluegrass (Poa
pratensis), timothy (Phleum pretense), smooth brome (Bro-
mus inermis), sedges (Carex spp.), western wheatgrass
(Pascopyrum smithii), prairie dropseed (Sporobolus heter-
olepis), fleabane (Erigeron spp.) and yarrow (Achillea spp.)
(Uresk etal. 2009). Additional ungulate species occupying
the study area included mule deer (Odocoileus hemionus),
white-tailed deer (O. virginianus), and elk (Cervus ela-
phus). Potential species known to predate on bighorn sheep
occurring in the study area included mountain lions [Puma
concolor (Smith etal. 2014; Wilckens etal. 2016; Jenks
2018)], coyotes (Canis latrans), bobcats [Lynx rufus (Parr
etal. 2014)], bald eagles (Haliaeetus leucocephalus), and
golden eagles (Aquila chrysaetos).
Translocation anddata collection
In February 2015, we (i.e., SDGFP and South Dakota State
University personnel) traveled to Hinton, Alberta, Canada
to capture and transport bighorn sheep to South Dakota.
T.J.Werdel et al.
1 3
Two sites within the Luscar Mine were baited with alfalfa
(Medicago sativa) hay for one week prior to our capture
date. On 10 February 2015, modified 18m × 18m elec-
tromagnetic drop-nets were constructed over bait sites,
and the following morning nets were used to capture 26
bighorn sheep (Jedrzejewski and Kamler 2004). Rams
were aged based on horn annuli (Geist 1966) and ewes
were aged via tooth eruption and wear (Hemming 1969;
Krausman and Bowyer 2003). Twenty-one adult ewes and
1 ram were fitted with store-on-board global positioning
Fig. 1 Locations of Black Hills National Forest (BHNF) domestic
sheep [South Dakota Game, Fish and Parks (SDGFP) survey 2015],
bighorn sheep range (SDGFP GPS and VHF collar data), city bound-
aries of Deadwood, South Dakota and Lead, South Dakota, and trans-
located Deadwood bighorn sheep study area, located in the Black
Hills of South Dakota, USA ( Modified from Werdel etal. 2020)
Space use andmovement patterns oftranslocated bighorn sheep
1 3
system (GPS; Model 2110D; 154–155MHz) and very high
frequency (VHF) radio collars (Advanced Telemetry Sys-
tems, Isanti, MN, USA) with a fix rate of 1 location every
5h (Fix Rate Success = 96.91%).
Translocated bighorn sheep were released approxi-
mately 3.5km southwest of Deadwood, South Dakota
on private land. All radio-collared bighorn sheep were
located (recorded with a hand-held GPS device [Garmin
Inc., Olathe, KS, USA]) a minimum of 5 times per week
post-release between February 2015–August 2015 and May
2016–July 2016, and a minimum of 3–4 times per week
September 2015–April 2016 and August 2016–December
2016 using a hand-held directional antenna and portable
receiver (model RA-23K, Telonics, Inc., Mesa, AZ, USA).
GPS collars fitted on live bighorn sheep released automati-
cally after ~ 700days and were recovered in the field, while
GPS collars fitted on bighorn sheep mortalities were recov-
ered at mortality sites; satellite location and time data were
offloaded from GPS collars into ArcMap 10.3.1 (Environ-
mental Systems Research Institute, Redlands, California,
USA). Ewes (all adult ewes were assumed pregnant) were
located (VHF) and observed [spotting scope from > 500m
(Karsh etal. 2016)] daily throughout the lambing period
(May–June) to monitor for evidence of parturition; once
evidence [observation of birth or lamb estimated 24h old
in bed site (Smith etal. 2015)] was obtained, the location
was estimated with a hand-held GPS device (Garmin Inc.,
Olathe, KS, USA), and later assessed in ArcGIS 10.3.1
(Environmental Systems Research Institute, Redlands,
California, USA). Slope (degrees) was estimated in Arc-
GIS 10.3.1 (Environmental Systems Research Institute,
Redlands, California, USA) utilizing Digital Elevation
Models (United States Geological Survey, The National
Map Viewer).
Data analysis
GPS locations [no locations were screened as 57,026/57,026
locations < 10 horizontal dilution of precision (Lowrey etal.
2017)] for 22 translocated bighorn sheep were transformed
to a spatial points data frame in Program R (R Core Team
2020) [“adehabitatHR” Package (Calenge 2006)]. The first
week of location data was removed to ensure movements
were representative of exploration behavior and habitat
attributes, and not a result of stress induced by manipu-
lations. Kernel density estimates (KDE) for home-ranges
were calculated at 99%, 95%, and 50% densities with
smoothing factors estimated with maximum log-likelihood
for each bighorn sheep individual (Worton 1989; Calenge
2006; Zimmerman 2008; R Core Team 2020). Minimum
convex polygons (MCP) (99%, 95%, and 50%) were cre-
ated using the same spatial points data frame in Program
R (R Core Team 2020) [“adehabitatHR” Package (Calenge
2006)]. We calculated 99%, 95%, and 50% Brownian Bridge
Movement Model (BBMM) home range contours using the
“BBMM” package in Program R (Nielson etal. 2017; Parr
2015; R Core Team 2020). We estimated home-range at
multiple densities (99%, 95%, and 50%) to compare effects
of initial pioneering behavior at differing home-range den-
sity scales.
Contour polygons for each method (KDE, MCP, and
BBMM) and density (99%, 95%, and 50%) were converted
to shapefiles (R Core Team 2020) and mapped in ArcGIS
10.3.1 (Environmental Systems Research Institute, Red-
lands, California, USA) to evaluate locations and home
ranges for each individual bighorn sheep. Individual home-
ranges for each method and density were combined using
the “union” tool in ArcGIS 10.3.1 (Environmental Systems
Research Institute, Redlands, California, USA) to deter-
mine the area utilized as home-range by the Deadwood
bighorn sheep herd. We isolated year 1 (Feb 2015–Jan
2016) and year 2 (Feb 2016–Jan 2017) locations for indi-
vidual bighorn sheep home-ranges to compare between
time periods.
Movement for each individual bighorn sheep, isolating
only locations obtained from year 1 (Feb 2015–Jan 2016)
and year 2 (Feb 2016–Jan 2017) time periods, were plot-
ted using the “move” package in Program R (Werdel etal.
2018; R Core Team 2020); movement statistics (i.e., travel
distance, maximum distance, minimum distance) also were
derived. Visual and statistical [travel distance (ANOVA)]
comparisons were made between the separate time periods
to test the hypothesis that translocated Deadwood bighorn
sheep herd would pioneer a greater distance (i.e., utilize
exploratory movements in excess of core home-range) in
year 1 than year 2.
Results
One ewe died due to capture stress shortly after release
(6 March 2015), resulting in too few GPS locations to be
utilized in analyses. Two subadults that were fitted with
expandable lamb collars were chemically immobilized and
lamb collars were replaced with new GPS store-on-board
adult collars (14 October 2015, 4 April 2016). Collars
released ~ 15 January 2017, and 22 of 23 were successfully
retrieved; one collar was left in the field as it was irretriev-
able, located on the side a high cliff face.
Immediately post-release (12 February 2015), translo-
cated bighorn sheep dispersed indiscriminately from the
release site. Through the first 2–3weeks, separate individual
bighorn sheep were observed at distances of 16km South,
18km West, 17km North, and 12km East of the release
T.J.Werdel et al.
1 3
site. After analyzing the location data in ArcGIS 10.3.1, it
was found that 1 ewe traveled 62km South before return-
ing back North to the release site. This pioneering behavior
by multiple individual bighorn sheep (individual locations
observed:
x
= 2592.09) continued for 3–4months post-
release until the lambing period (May–June) began. During
the lambing period, pregnant ewes sought out rugged, iso-
lated areas. Two lambing areas were identified in the field
and confirmed by isolating GPS locations during the lamb-
ing period (May–June 2015 and 2016) uploaded into ArcGIS
10.3.1. These lambing areas (Fig.1) were utilized in both
May–June 2015 and May–June 2016 and had significantly
[F(df(1, 57,024))] = 875.58, p = < 0.01) higher slope values
(
x
= 27.36 degrees, n = 1486, SD = 11.78) than locations
within the entire study area (
x
= 20.58 degrees, n = 55,540,
SD = 8.62). Home range sizes varied among methods [KDE
(99%, 95%, 50%; Fig.2), MCP (99%, 95%, 50%; Fig.3),
and BBMM (99%, 95%, 50%; Figs.4, 5, 6) (Table1)], but
estimates were not statistically significant. KDE, MCP,
and BBMM mean individual mean home-range sizes, and
union of home-range sizes were significantly larger in year
1 (n = 22), than year 2 (n = 19) (excluding BBMM 50%)
(Tables2 and 3).
Only 4 GPS collars were recovered from bighorn sheep
that survived the entire study period (Feb 2015–Jan 2017)
(19 of 23 originally collared (GPS) bighorn sheep died
during the study period; recruitment = 40, mortality = 44,
population as of Jan 2017 = 24), which allowed compari-
sons between year 1 and year 2 movement data. BH223
(bighorn sheep identification number) traveled a total
distance of 339.68km in year 1 and 384.44km in year 2
(Fig.7a), resulting in 46.9% of total travel occurring in
year 1. BH232 traveled a total distance of 444.46km in
year 1 and 318.72km in year 2 (Fig.7b), resulting in 58.2%
of total travel occurring in year 1. BH233 traveled a total
distance of 600.27km in year 1 and 340.07km in year 2
(Fig.7c), resulting in 63.8% of total travel occurring in year
1. BH239 traveled a total distance of 342.78km in year 1
and 431.85km in year 2 (Fig.7d), resulting in 44.3% of total
travel occurring in year 1.
Discussion
As we predicted, translocated bighorn sheep in our study
exhibited similar post-release behavior to ungulate species
translocated worldwide; pioneering a novel environment
(e.g., relatively large distances traversed and large home
ranges) for a temporary duration (e.g., year 1) before the
establishment of a stable home range (e.g., year 2) [Bleich
etal. 1996 (Ovis canadensis nelsoni: USA); Kissell 1996
(Ovis canadensis: USA); Singer etal. 2000a, b (Ovis
canadensis: USA); Schmitz etal. 2015 (Bison bonasus:
Germany); Bleisch etal. 2017 (Cervus elaphus: USA);
Mertes etal. 2019 (Oryx dammah: Republic of Chad)]. To
accurately define a home range for translocated ungulates,
it may be appropriate to exclude pioneering data points; for
mountain ungulates (i.e., bighorn sheep) that rely on escape
terrain and movement corridors, it may also be beneficial
to utilize a home range estimator, such as BBMM, that uses
movement trajectories between locations. Due to the unpre-
dictability of space-use post-release of translocated ungu-
lates, we believe the BBMM home range estimator is most
appropriate for delineating areas, which not only exclude
early pioneering locations (low time duration between suc-
cessive points), but also accurately describe areas where
the species conducts its “normal activities of food gather-
ing, mating, and caring for young (Burt 1943).” BBMM
was shown to encompass similar areas as MCP and KDE,
but excluded areas of little or no observed use by bighorn
sheep.
Our 50% and 95% KDE mean home-range sizes (4.77 km2
and 41.41 km2) were much smaller in the area when com-
pared to KDE home-range sizes (14.38 km2 and 94.30 km2)
of Parr (2015), who conducted a similar study on bighorn
sheep in the southern Black Hills. However, Zimmerman
(2008) utilized 95% adaptive and fixed kernel home-range
analyses in her Badlands National Park (South Dakota) big-
horn sheep study, finding mean home-range sizes of 15.5
km2 (Adaptive kernel) and 16.1 km2. Zimmerman (2008)
also analyzed 50% adaptive (mean = 3.1 km2) and fixed ker-
nel (mean = 1.8 km2) home-range. Our 50% and 95% kernel
density mean home-range sizes fall between results from
these 2 similar South Dakota studies.
Previous studies on bighorn sheep utilizing MCP to
determine the home-range size (Leslie and Douglas 1979;
Seegmiller and Ohmart 1981; Krausman etal. 1989; Zim-
merman 2008) resulted in mean home-range sizes (mean
of summer, winter, male, and female home-ranges) of
5.65 km2, 12.30 km2, 23.48 km2, and 16.30 km2, respec-
tively, which were much smaller than our overall mean
95% MCP home-range size of 55.73 km2. However, when
we isolated only year 2 locations, our year 2 mean 95%
MCP home-range size of 12.91 km2 was similar to the
findings of Leslie and Douglas (1979), Seegmiller and
Ohmart (1981), Krausman etal. (1989), and Zimmerman
(2008). It is important to note that our MCP analysis
included areas not observed in the field as areas of use
by bighorn sheep.
When comparing BBMM of Parr’s (2015) study and this
study, we see similar results; Parr’s (2015) 95% BBMM
ranged from 15.68 km2 to 42.06 km2, while our 95% BBMM
Space use andmovement patterns oftranslocated bighorn sheep
1 3
mean home-range size was very similar (32.95 km2). As
Parr (2015) indicated, their limited location data may have
overestimated true home-range size at the larger extremes
(Horne 2007; Sawyer etal. 2009; Walter etal. 2011). Our
median home-range size may be due to large numbers of
GPS locations (n = 57,026), origin of the translocated big-
horn sheep, population sizes, or differing terrain and habitat
capabilities of the 2 study areas.
The Luscar Mine capture site, although similar in
topography to the release site, differed in population
Fig. 2 Combined home-ranges for 99% (678.39 km2), 95% (268.56 km2), and 50% (28.17 km2) Kernel Density Estimation (KDE) home-range
analyses of the translocated the Deadwood bighorn sheep herd in the Black Hills, South Dakota, USA (BHNF is Black Hills National Forest)
T.J.Werdel et al.
1 3
structure [~ 1000 individuals (Teck 2012), with specific
herds including as many as 193 individuals (47% ewes,
29% rams, and 24% lambs; MacCallum and Geist 1992)].
MacCallum and Geist (1992) found that bighorn sheep
they surveyed over multiple seasons (Pre-rut, Rut, Win-
ter, Spring Lambing, and Summer) utilized a 95% MCP
of 4.79 km2. This is much smaller than our overall mean
95% MCP (55.73 km2), but similar to our year 2 95%
Fig. 3 Combined home-ranges for 99% (788.95 km2), 95% (366.87 km2), and 50% (19.78 km2) Minimum Convex Polygon (MCP) home-range
analyses of the translocated Deadwood bighorn sheep herd in the Black Hills, South Dakota, USA (BHNF is Black Hills National Forest)
Space use andmovement patterns oftranslocated bighorn sheep
1 3
MCP (12.91 km2). Therefore, after acclimation to the
Black Hills region, the Deadwood bighorn sheep herd
may utilize habitat in a similar way as do their source
populations.
We utilized and compared 3 different home-range models
(KDE, MCP, and BBMM), finding vastly different areas
with each. However, after isolating year 2 locations, 95%
KDE (
x
= 9.83 km2), MCP (
x
= 12.91 km2), and BBMM
Fig. 4 Combined home-ranges for 99% (446.62 km2), 95% (102.83
km2), and 50% (1.18 km2) Brownian Bridge Movement Model
(BBMME) home-range analyses of the translocated Deadwood big-
horn sheep herd in the Black Hills, South Dakota, USA (BHNF is
Black Hills National Forest)
T.J.Werdel et al.
1 3
(
x
= 4.08 km2) (Fig.8) means were much more similar than
for the entire study period. Having only 4 originally fitted
GPS collars record data for the entire study period yielded
a relatively small sample size when comparing movement
from year 1 to year 2. However, the plotted movement mod-
els created with the “move” package in Program R allowed
us to visually examine distances and movement patterns of
the 4 individuals. The total distance traveled for each time
Fig. 5 Combined home-ranges for Year 1 (Feb 2015–Jan 2016)
99% (569.26 km2), 95% (187.33 km2), and 50% (1.72 km2) Brown-
ian Bridge Movement Model (BBMM) home-range analyses of the
translocated Deadwood bighorn sheep herd in the Black Hills, South
Dakota, USA (BHNF is Black Hills National Forest)
Space use andmovement patterns oftranslocated bighorn sheep
1 3
period may not be significant statistically, but when view-
ing each set of plots there were striking differences. For
each of the 4 sets of plots, year 1 showed more excursions
away from a centrally located mass of location points than
did year 2.
Established bighorn sheep herds may be migratory,
occupying multiple (3–6) distinct home-ranges through-
out the year (Geist 1971; Festa-Bianchet 1988; Parr
2015). The translocated Deadwood bighorn sheep herd
Fig. 6 Combined home-ranges for Year 2 (Feb 2016–Jan 2017) 99%
(56.78 km2), 95% (14.22 km2), and 50% (0.58 km2) Brownian Bridge
Movement Model (BBMM) home-range analyses of the translocated
Deadwood bighorn sheep herd in the Black Hills, South Dakota, USA
(BHNF is Black Hills National Forest)
T.J.Werdel et al.
1 3
was not observed as migratory. However, the only adult
male bighorn sheep that was originally translocated did
show signs of differing movement during the summer, but
was euthanized, due to potential contact with domestic
sheep, ending data collection (Werdel etal. 2018); this
type of movement by adult rams has been documented by
DeCesare and Pletscher (2006) and Parr (2015). Seasonal
movement was observed during the lambing season, which
was predicted due to previous bighorn sheep research in
the southern Black Hills (Parr 2015). During the lamb-
ing period, ewes tend to utilize habitat with less nutri-
tious areas that include cliffs and large amounts of escape
terrain than throughout the remainder of the year (Geist
1971; Leslie and Douglas 1979).
Table 1 Results of Kernel Density Estimation (KDE), Minimum Convex Polygon (MCP), and Brownian Bridge Movement Model (BBMM)
home-range estimators on GPS collared Deadwood bighorn sheep (n = 22)
Included in the table are mean home-range sizes (
x
; km2), standard error of mean home-range sizes (SE), and area encompassed by the union of
individual home-ranges (∪) utilizing each of the 3 methods discussed at 50%, 95%, and 99% estimates
50% 95% 99%
x
SE
x
SE
x
SE
Kernel Density 4.77 1.06 28.17 41.41 10.50 268.56 91.22 26.40 678.39
MCP 3.24 0.84 19.78 55.73 15.04 366.87 131.93 35.05 788.946
BBMM 1.57 1.08 1.18 32.95 4.67 102.83 60.41 7.76 446.62
Table 2 Comparisons between
year 1 (Feb 2015–Jan 2016)
and year 2 (Feb 2016–Jan
2017) Kernel Density
Estimation (KDE), Minimum
Convex Polygon (MCP) and
Brownian Bridge Movement
Model (BBMM) home-range
estimators on GPS collared
Deadwood bighorn sheep
(n = 22)
Included in the table are mean home-range sizes (
x
; km2), standard error of mean home-range sizes (SE),
and area encompassed by the union of individual home-ranges () utilizing each of the 3 methods dis-
cussed at 50%, 95%, and 99% estimates
50% 95% 99%
x
SE
x
SE
x
SE
Kernel Density Year 1 7.29 1.87 50.45 65.34 24.21 573.14 124.64 46.19 1066.06
Year 2 1.51 0.16 5.88 9.83 1.09 34.95 20.12 2.39 71.82
MCP Year 1 5.11 1.12 27.44 65.01 18.18 405.34 131.93 35.05 1175.06
Year 2 1.68 0.22 5.36 12.91 3.19 78.64 29.05 3.73 93.18
BBMM Year 1 1.57 1.08 1.72 40.01 5.61 187.33 58.06 8.61 569.26
Year 2 0.27 0.07 0.58 4.08 0.29 14.22 11.04 1.34 56.78
Table 3 Comparisons between year 1 (Feb 2015–Jan 2016) and year 2 (Feb 2016–Jan 2017) kernel density estimation (KDE), minimum convex
polygon (MCP) and brownian bridge movement model (BBMM) home-range estimators on GPS collared deadwood bighorn sheep (n = 22)
Included in the table are F statistic values [F (degrees of freedom)] and P values (P) of ANOVA calculations between year 1 and year 2 indi-
vidual home-ranges, utilizing each of the 3 methods discussed at 50%, 95%, and 99% estimates
50% 95% 99%
F(1, 39)
P
F(1, 39)
P
F(1, 39)
P
Kernel density 8.18 0.01 4.52 0.04 4.29 0.05
MCP 7.86 0.01 6.91 0.01 7.35 0.01
BBMM 0.01 0.97 35.24 < 0.01 25.220 < 0.01
Space use andmovement patterns oftranslocated bighorn sheep
1 3
Conclusions andmanagement implications
Currently (Jan 2021) the Deadwood bighorn sheep herd is
occupying the original study area. The Deadwood bighorn
sheep herd may require a smaller home-range size than do
other bighorn sheep herds in the Black Hills, possibly due to
the smaller home-range sizes of Luscar Mine source popu-
lations. Immediate pioneering post-release of translocated
bighorn sheep should be expected in future reintroductions
to novel habitats and may contribute to relatively high mor-
tality rates due to disease transmission from domestic sheep
(Werdel etal. 2020). Further augmentation of the Deadwood
bighorn sheep herd may be required; the potential for pio-
neering would likely be reduced with an established herd
already occupying the area. The lambing areas outlined in
Fig.1 will most likely continue to be utilized by the Dead-
wood bighorn sheep herd on a yearly/seasonal basis. As
much of the home range is near or overlapping major roads
and communities, it would be beneficial to managers to
campaign for slower speed limits and increase the visibil-
ity/frequency of bighorn sheep crossing signs. We recom-
mend continued monitoring of the Deadwood bighorn sheep
herd to assess whether movement patterns and home-ranges
remain the same, or if seasonal migration begins to occur. It
may also benefit managers to begin evaluating home-range
size and movements after pioneering behavior slows or
ceases; beginning year 2 post-release of translocated big-
horn sheep.
Fig. 7 Movement comparison of Year 1 (Black) and Year 2 [Feb
2016–Jan 2017 (Grey)] of 4 translocated Deadwood bighorn sheep in
the Black Hills, South Dakota, USA, via the “move” package in Pro-
gram R (UTM Zone 13). BHnumbers(e.g., BH223, BH232, BH233,
BH239) indicate individual bighorn sheep identification numbers and
letters (a, b, c, d) are subfigures referenced in the main text
T.J.Werdel et al.
1 3
Acknowledgements We thank the South Dakota Department of Game,
Fish and Parks, Civil Air Patrol, Deadwood Police Department, Law-
rence County Sheriff’s Office, and private property owners in the
Deadwood area for their assistance and property access. We thank J.
Smith and B. Simpson for their assistance with data analyses. We thank
T. Haffley, K. Cudmore, J. Doyle, J. Clark, and C. Werdel for their
assistance with monitoring, capturing, and euthanizing bighorn sheep
during the study period.
Author contributions TJW, JAJ, JTK, CPL conceived and designed
the study; TJW conducted field data collection; JTK led reintroduction
efforts; TJW conducted data analyses; TJW wrote the manuscript and
Fig. 8 Comparison of combined Year 2 (Feb 2016–Jan 2017) 95%
Brownian Bridge Movement Model (BBMM) home-range (14.22
km2), Year 2 95% Kernel Density Estimator (KDE) home-range
(34.95 km2), and Year 2 95% Minimum Convex Polygon (MCP)
home-range (78.64 km2) analyses of the translocated Deadwood big-
horn sheep herd in the Black Hills, South Dakota, USA (BHNF is
Black Hills National Forest)
Space use andmovement patterns oftranslocated bighorn sheep
1 3
all authors discussed results and contributed to the final manuscript;
JAJ secured funding.
Funding Financial support for this project was provided by Federal
Aid to Wildlife Restoration administered through the South Dakota
Department of Game, Fish and Parks (Study Number 7556).
Data availability The data used and/or analyzed during the study are
available from the corresponding author with a reasonable request.
Compliance with ethical standards
Conflict of interest The authors declare that they have no competing
interests.
Ethics approval and consent to participate All capture and handling
methods were approved by the South Dakota State University Insti-
tutional Animal Care and Use Committee (Approval No. 14-096A).
Access to private property, when needed, was done so with landowner
permission. No threatened or protected species were involved in this
research.
References
Barg JJ, Jones J, Robertson RJ (2005) Describing breeding territories
of migratory passerines: suggestions for sampling, choice of esti-
mator, and delineation of core areas. J Anim Ecol 74:139–149
Beecham JJ, Collins CP, Reynolds TD (2007) Rocky mountain bighorn
sheep (Ovis canadensis): a technical conservation assessment.
USDA Forest Service, Rocky Mountain Region
Bleich VC, Wehausen JD, Ramey RR, Rechel JL (1996) Metapopula-
tion theory and mountain sheep: implications for conservation. In:
McCullough DR (ed) Metapopulations and wildlife conservation.
D.C., Island Press, Washington, pp 353–373
Bleich VC, Bowyer RT, Wehausen JD (1997) Sexual segregation in
mountain sheep: resources or predation? Wildlife Monographs
134:3–50
Bleisch AD, Keller BJ, Bonnot TW, Hansen LP, Millspaugh JJ (2017)
Initial movements of reintroduced Elk in the Missouri Ozarks.
Am Midl Nat 178:1–16
Borger L, Franconi N, De Michele G, Gantz A, Meschi F, Manica A,
Lovari S, Coulson T (2006) Effects of sampling regime on the
mean and variance of home range size estimates. J Animal Ecol
75:1393–1405
Buechner HK (1960) The bighorn sheep in the United States, its past,
present, and future. Wildlife Monogr 4:3–174
Burt WH (1943) Territoriality and home range concepts as applied to
mammals. J Mammal 24:346–352
Calenge C (2006) The package adehabitat for the R software: tool
for the analysis of space and habitat use by animals. Ecol Model
197:1035
DeCesare NJ, Pletscher DH (2006) Movements, connectivity, and
resource selection of Rocky Mountain bighorn sheep. J Mammal
87:531–538
Douglas CL, Leslie DM Jr (1999) Management of bighorn sheep. In:
Valdez R, Krausman PR (eds) Mountain sheep of North Amer-
ica. Arizona, USA, The University of Arizona Press, Tucson, pp
238–262
Festa-Bianchet M (1986) Seasonal dispersion of overlapping mountain
sheep ewe groups. J Wildlife Manag 50:325–330
Festa-Bianchet M (1988) Seasonal range selection in bighorn sheep:
conflicts between forage quality, forage quantity, and predator
avoidance. Oecologia 75:580–586
Geist V (1966) Validity of horn segment counts in aging bighorn sheep.
J Wildl Manag 30:634–635
Geist V (1971) Mountain sheep: a study in behavior and evolution.
Illinois, USA, University of Chicago Press, Chicago
Getz WM, Wilmers CC (2004) A local nearest-neighbor convex-hull
construction of home ranges and utilization distributions. Ecog-
raphy 27:489–505
Gogan PJ (1990) Considerations in the reintroduction of native
mammalian species to restore natural ecosystems. Nat Areas J
10:210–217
Gordon IG, Hester AJ, Festa-Bianchet M (2004) The management of
wild large herbivores to meet economic, conservation and envi-
ronmental objectives. J Appl Ecol 41:1021–1031
Griffith B, Scott JM, Carpenter JW, Reed C (1989) Translocation
as a species conservation tool: status and strategy. Science
245:477–480
Hemming JE (1969) Cemental deposition, tooth succession, and horn
development as criteria of age in dall sheep. J Wildl Manag
33:552–558
Hoffman GR, Alexander RR (1960) Forest vegetation of the black hills
national forest of south dakota and wyoming: a habitat type clas-
sification. USDA research paper RM-276: rocky mountain forest
and range experiment station, Fort Collins, Colorado, USA 48
Horne JS, Garton EO, Krone SM, Lewis JS (2007) Analyzing animal
movements using Brownian bridges. Ecology 88:2354–2363
Jacques CN, Jenks JA, Deperno CS, Sievers JD, Grovenburg TW,
Brinkman TJ (2009) Evaluating ungulate mortality associated
with helicopter net-gun captures in the Northern Great Plains. J
Wildlife Manag 73:1282–1291
Jedrzejewski W, Kamler JF (2004) Modified drop-net for capturing
ungulates. Wildl Soc Bull 32:1305–1308
Jenks JA (2018) Mountain lions of the Black Hills: history and ecology.
Johns Hopkins University Press, Baltimore, Maryland
Karsch RC, Cain JW, Rominger EM, Goldstein EJ (2016) Desert big-
horn sheep lambing habitat: parturition, nursery, and predation
sites. J Wildl Manag 80:1069–1080
Kie JG, Matthiopoulos J, Fieberg J, Powell RA, Cagnacci F, Mitchell
MS (2010) The home-range concept: are traditional estimators
still relevant with modern telemetry technology? Phil Trans R
Soc B 365:2221–2231
Kissell RE (1996) Competitive interactions among bighorn sheep,
feral horses and mule deer in Bighorn Canyon National Rec-
reation Area. Ph.D. Dissertation. Bozeman, MT, Montana State
University
Krausman PR, Bowyer RT (2003) Mountain sheep (Ovis canadensis
and O. dalli). In: Feldhamer GA, Thompson BC, Chapman JA
(eds) Wild mammals of North America: biology, management,
and conservation, 2nd edn. JHU Press, Baltimore, Maryland, pp
1095–1115
Krausman PR, Cox AS (2019) Vexing vocabulary in submissions to the
Journal of Wildlife Management. J Wildl Manag 83:1279–1280
Krausman PR, Leopold BD, Seegmiller RF, Torres SG (1989) Rela-
tionships between desert bighorn sheep and habitat in western
Arizona. Wildl Monogr 102:3–66
Laver PN, Kelly MJ (2008) A critical review of home range studies.
Journal of Wildlife Management 72:290–298
Laver PN, Kelly MJ (2010) A critical review of home range studies. J
Wildl Manag 72:290–298
Leslie DM, Douglas CL (1979) Desert bighorn sheep of the River
Mountains, Nevada. Wildl Monogr 66:3–56
Lewis JS (2007) The effects of human influences on black bear habitat
selection and movement patterns within a highway corridor. Uni-
versity of Idaho, Moscow, Thesis
T.J.Werdel et al.
1 3
Lowrey B, Garrott RA, Miyasaki HM, Fralick G, Dewey SR (2017)
Seasonal resource selection by introduced mountain goats in the
southwest Greater Yellowstone Area. Ecosphere 8(4):e01769.
https:// doi. org/ 10. 1002/ ecs2. 1769
MacCallum NB, Geist V (1992) Mountain restoration: soil and surface
wildlife habitat. Geo J 27:23–46
Maccallum NB (2012) Effects of delayed spring greenup on bighorn
sheep of the Luscar and Gregg River mines, Alberta. Bienn Symp
North Wild Sheep Goat Council 18:71–78
Mcintosh AC (1949) A botanical survey of the Black Hills of South
Dakota. Black Hills Eng 28:1–74
Mertes K, Stabach JA, Songer M, Wacher T, Newby J, Chuven J, Al
Dhaheri S, Leimgruber P, Monfort S (2019) Management back-
ground and release conditions structure post-release movements
in reintroduced ungulates. Front Ecol Evolut 7:1–14
National oceanic and atmospheric administration (2017) www. ncdc.
noaa. gov. Accessed 10 Jan 2017
Nicholson MC, Bowyer RT, Kie JG (1997) Habitat selection and sur-
vival of mule deer: tradeoffs associated with migration. J Mammal
78:483–504
Nielson RM, Sawyer H, McDonald TL (2017) BBMM: Brownian
bridge movement models. 2013. http:// CRAN.R- proje ct. org/ packa
ge= BBMM. Accessed Jan 2017
Nilsen EB, Pedersen S, Linnell JDC (2008) Can minimum convex
polygon home ranges be used to draw biologically meaningful
conclusions? Ecol Res 23:635–639
Orr HK (1959) Precipitation and streamflow in the Black Hills. Station
Paper RM44. U.S. Department of Agriculture, Forest Service,
Rocky Mountain Forest and Range Experiment Station, Fort Col-
lins, CO. p. 25
Parr BL, Kanta J, Sandrini J, Thompson DJ, Jenks JA (2014) Bobcat
predation on bighorn lamb in the western Black Hills of South
Dakota. Prairie Nat 46:41–43
Parr BL (2015) Population parameters of a bighorn sheep herd inhabit-
ing the Elk Mountain region of South Dakota and Wyoming. M.S.
Thesis. Brookings, SD, South Dakota State University 148
Powell RA, Mitchell MS (2012) What is a home range? J Mammal
93:948–958
Powell RA (2000) Animal home ranges and territories and home range
estimators. In: Pearl MC (ed) Research techniques in animal ecol-
ogy. Columbia University Press, New York, pp 65–103
R Core Team (2020) R: a language and environment for statistical com-
puting. R Foundation for Statistical Computing, Vienna, Austria
Richardson AH, Peterson LE (1974) History and management of South
Dakota deer. South Dakota Department of Game, Fish and Parks,
Pierre, South Dakota, Bulletin 5: p. 113.
Sawyer H, Kauffman MJ, Nielson RM, Horne JS (2009) Identifying
and prioritizing ungulate migration routes for landscape-level
conservation. Ecol Appl 19:2016–2025
Schmitz P, Caspers S, Warren P, Witte K (2015) First steps into the
wild - exploration behavior of European bison after the first rein-
troduction in Western Europe. PLoS ONE 10:e0143046
Schuler KL, Schroeder GM, Jenks JA, Kie JG (2014) Ad hoc smooth-
ing parameter performance in kernel estimates of GPS-derived
home ranges. Wbio 20(5):259–266
Scillitani L, Darmon G, Monaco A, Cocca G, Sturaro E, Rossi L,
Ramanzin M (2013) Habitat selection in translocated gregarious
ungulate species: an interplay between sociality and ecological
requirements. J Wildl Manag 77:761–769
Seegmiller RF, Ohmart RD (1981) Ecological relationships of feral
burros and desert bighorn sheep. Wildl Monogr 78:3–58
Seton ET (1929) Lives of game animals. Doubleday, Doran and Co.,
Inc., New York, NY 4 441–501
Singer FJ, Bleich VC, Gudorf MA (2000b) Restoration of bighorn
sheep metapopulations in and near western national parks. Restor
Ecol 8:14–24
Singer FJ, Papouchis CM, Symonds KK (2000a) Translocations as
a tool for restoring populations of bighorn sheep. Restor Ecol
8:6–13
Smith JB, Grovenburg TW, Jenks JA (2015) Parturition and bed site
selection of bighorn sheep at local and landscape scales. J Wildl
Manag 79:393–401
Smith JB, Jenks JA, Grovenburg TW, Klaver RW (2014) Disease and
predation: sorting out causes of a bighorn sheep (Ovis canadensis)
decline. PLoS ONE 9:e88271
South Dakota Department of Game, Fish and Parks. Action plan for
management of bighorn sheep in South Dakota (2013) http:// gfp.
sd. gov/ wildl ife/ manag ement/ plans. Accessed 27 February 2017
Teck (2012) Teck 2012 sustainability report. Teck Resources Limited,
Vancouver, British Columbia, Canada, p 148
Thilenius JF (1972) Classification of deer habitat in the ponderosa pine
forest of the Black Hills, South Dakota. USDA Forest Service
Research Paper RM-91. Fort Collins, Colorado 28
Uresk DW, Mergen DE, Benzon TA (2009) Monitoring meadows with
a modified Robel pole in the northern Black Hills, South Dakota.
Prairie Nat 41:121–125
Van Vuren D (1998) Mammalian dispersal and reserve design. In: Caro
T (ed) Behavioral ecology and conservation biology. NY, Oxford
University Press, New York, pp 369–393
Walter WD, Fischer JW, Baruch-Mordo S, VerCauteren KC (2011)
What is the proper method to delineate home range of an animal
using today’s advanced GPS telemetry systems: the initial step. In:
Krejcar O (ed) Modern telemetry, InTech, pp 249–268
Werdel TJ, Jenks JA, Besser TA, Kanta JT, Lehman CP, Frink TJ
(2018) Survival of translocated bighorn sheep in the Deadwood
region of the Black Hills, South Dakota. Northwestern Nat
99:222–231
Werdel TJ, Jenks JA, Besser TA, Kanta JT, Lehman CP, Frink TJ
(2020) Restoration of a bighorn sheep population impeded by
Mycoplasma ovipneumoniae exposure. Restor Ecol 28:387–395
Wilckens DT, Smith JB, Tucker SA, Thompson DJ, Jenks JA (2016)
Mountain lion (Puma concolor) feeding behavior in the Little
Missouri Badlands of North Dakota. J Mammal 97:373–385
Wilckens D (2014) Ecology of mountain lions (Puma concolor) in the
North Dakota Badlands: population dynamics and prey use. M.S.
Thesis. Brookings, SD, South Dakota State University 92
Witte SS, Gallagher MV (2012) The North American Journals of
Prince Maximilian of Wied, vol 3. Oklahoma, USA, University
of Oklahoma Press, Norman
Worton BJ (1987) A review of models of home range for animal move-
ment. Ecol Model 38:277–298
Worton BJ (1989) Kernel methods for estimating the utilization distri-
bution in home range studies. Ecology 70:164–168
Worton BJ (1995) A convex hull-based estimator of home-range size.
Biometrics 51:1206–1215
Zimmerman TJ (2008) Evaluation of an augmentation of Rocky Moun-
tain bighorn sheep at Badlands National Park, South Dakota.
Ph.D. Dissertation. Brookings, SD, South Dakota State Univer-
sity 139
Publisher’s Note Springer Nature remains neutral with regard to
jurisdictional claims in published maps and institutional affiliations.
... Despite substantial advancements within the last 15 years, up to 45% of recent translocation projects failed or were considered a "partial success" only (Soorae, 2011(Soorae, , 2013(Soorae, , 2016. One of the reasons for failure is that animals typically exhibit extensive movements following their release because of the need to explore their new habitat (Bleisch et al., 2017;Moehrenschlager & Macdonald, 2003;Werdel et al., 2021). These large movements are energetically costly (Bonte et al., 2012) and increase exposure to predators (Leech et al., 2017;Yoder et al., 2004) or the risk of accidents such as vehicle collisions (Spinola et al., 2008), ultimately leading to greater mortalities. ...
... In some countries where they are declining, several species have been subject to extensive reintroduction programs (e.g. kulan, Equus hemionus kulan, Werdel et al., 2021;woodland caribou, Rangifer tarandus caribou, Leech et al., 2017). However, it is not easy to predict translocation outcomes in ungulates since their post-release behaviour is influenced by several parameters including habitat quality (Smith et al., 2022), similarities between the source and release sites (Scillitani et al., 2013), age , sex (Bocci et al., 2016), release season (Garnier et al., 2021) and method (i.e. ...
Thesis
Full-text available
Translocations, “the deliberate movement of organisms from one site for release in another”, are increasingly being used for wildlife conservation and management. However, their success rate is still relatively low. Failures of translocation projects have often been attributed to the extensive movements made by newly-released individuals or their inability to acclimatise. However, it is unclear if animals display this typical movement pattern when released into fenced areas where movements are restricted. Furthermore, we do not know if methods commonly used to facilitate acclimatisation work in those systems. In particular, although the presence of conspecifics has sometimes shown to facilitate establishment, we do not know if this holds true for non-social ungulate species. Using GPS data on moose (Alces alces) and red deer (Cervus elaphus) released in a small fenced reserve, we aimed at 1) describing the post-release spatial behaviour of both species and identifying the time needed to acclimatise and 2) determining if the presence of conspecifics influences the length of the acclimatisation period in a non-social species. We found that neither moose nor red deer had larger home ranges in the first weeks post-release. Instead, both speciestended to have comparatively smaller home ranges during this period. Red deer had longer step lengths following release, but in the first 80 days only. Moose seemed to acclimatise immediately after translocation, whereas deer had an acclimatisation period of around 15 weeks. Home ranges and step lengths of the first moose cohort tended to be larger in the first weeks post-release only, suggesting some influence of conspecifics. These results show that moose and red deer seem to acclimatise relatively shortly after translocation, but limits imposed by the enclosure might play a role. Multitrait studies are needed to assess the full impact of confinement on post-release spatial behaviour to improve translocation outcomes.
... 이 외에도 연구자의 결정에 따라 95%나 85% MCP를 사용 하기도 하고 (Blondel et al., 2009), 50% MCP 행동권을 이용하여 핵심(core) 행동권을 도출하기도 한다 (Franzreb, 2006). 하지만, MCP 방법은 행동권의 크기를 과대 추정한 다는 점과 새로운 행동권 추정 방법론 개발로 인해 과거의 방법론으로 취급되고 있으며 (Getz and Wilmers, 2004;Kie et al., 2010), 최근 연구에선 대부분 새로운 방법론에 대한 비교 대상으로 사용되고 있다 (Franzreb, 2006;Holt et al., 2010;Werdel et al., 2021). ...
Article
Animals exhibit certain behaviors and movement patterns as they react to their internal needs, external stimuli, and surrounding environments. They have a bounded range in which they mostly spend their time, and it is referred to as a home range. Based on the fact that the home range is a critical area for the survival and preservation of species, there has been a growing body of research on developing more precise home range estimation methods to use the estimated ranges as a ground for establishing an effective conservation policy since the early 1940s. Recent rapid advancements in telemetry technology that resulted in the presence of autocorrelation between locations with short time intervals revealed the limitations of the existing estimators. Many novel estimators have been developed to compensate for it by incorporating autocorrelation in calculating home ranges. However, studies on the animal home range are still in their early stage in Korea, and newly developed methodologies have not yet been adopted. Therefore, this study aims to introduce the foreign home range estimation methods and foster domestic research activities on home ranges. Firstly, we compared and contemplated seven estimators by categorizing them into geometrical and statistical methodologies and then divided them into estimators that assume independent observations and those that consider autocorrelation in each category. After that, the home ranges of black-tailed gulls (Larus crassirostris) were calculated using GPS tracking data for the month of June and derived home range estimators by applying the methodology introduced in this study. We analyzed and compared the results to discuss the strengths and weaknesses of each method. Lastly, we proposed a guideline that can help researchers choose an appropriate estimator for home range calculation based on the animal location data characteristics and analysis purpose.
... An animal's habitat selection changes depending on current motivation and internal state (Nathan et al., 2008;Roever et al., 2014), and motivation is fundamentally different when animals are focused on exploration vs. exploitation. This issue is frequently ignored, or sometimes dealt with by discarding data for the first few days or weeks after release (e.g., Mondal et al., 2013;Werdel et al., 2021), during which behavior is assumed not to be representative of a steady state. In the latter case, the choice of the temporal cut-off to use is arbitrary and relies on the assumption that every individual in the population behaves similarly. ...
Article
Full-text available
Translocated animals undergo a phase of behavioral adjustment after being released in a novel environment, initially prioritizing exploration and gradually shifting toward resource exploitation. This transition has been termed post-release behavioral modification. Post-release behavioral modification may also manifest as changes in habitat selection through time, and these temporal dynamics may differ between individuals. We aimed to evaluate how post-release behavioral modification is reflected in temporal dynamics of habitat selection and its variability across individuals using a population of translocated female greater sage-grouse as a case study. Sage-grouse were translocated from Wyoming to North Dakota (USA) during the summers of 2018–2020. We analyzed individual habitat selection as a function of sagebrush cover, herbaceous cover, slope, and distance to roads. Herbaceous cover is a key foraging resource for sage-grouse during summer; thus, we expected a shift from exploration to exploitation to manifest as temporally-varying selection for herbaceous cover. For each individual sage-grouse ( N = 26), we tested two competing models: a null model with no time-dependence and a model with time-dependent selection for herbaceous cover. We performed model selection at the individual level using an information-theoretic approach. Time-dependence was supported for five individuals, unsupported for seven, and the two models were indistinguishable based on AIC c for the remaining fourteen. We found no association between the top-ranked model and individual reproductive status (brood-rearing or not). We showed that temporal dynamics of post-release habitat selection may emerge in some individuals but not in others, and that failing to account for time-dependence may hinder the detection of steady-state habitat selection patterns. These findings demonstrate the need to consider both temporal dynamics and individual variability in habitat selection when conducting post-release monitoring to inform translocation protocols.
Article
Full-text available
One of the greatest challenges in restoring species to the wild is insufficient knowledge about their habitat requirements and movement ecology. This is especially true for wide-ranging species such as the scimitar-horned oryx (Oryx dammah). Once widespread across Sahelo-Saharan grasslands, oryx were declared Extinct in the Wild in 1999. Here, we integrate GPS/satellite tracking, remote sensing, and movement analyses to assess how reintroduced oryx respond to wild conditions. We monitored two groups of oryx, reared under different captive management regimes and released in different seasons, for 12 months after release. Our study provides the first movement trajectories and home range estimates for this species. We expected oryx movements after release to represent trade-offs between risky, energetically expensive exploration and resource exploitation. Oryx raised under semi-free ranging conditions and released during the wet season ("ranging") exhibited this pattern of exploration followed by home range establishment. In contrast, oryx raised in small pens and released during the dry season ("penned") explored far less novel terrain. Ranging oryx exhibited seasonal shifts in activity and movement timing, while penned oryx simply reduced overall movement and continuously accessed supplemental food and water. Sahelian ecosystems exhibit strong seasonal cycles and extensive spatial variation. In this highly variable environment, reintroduced oryx will need to disperse from the release site to acquire adequate forage throughout the year. Thus, we experimentally varied acclimation period, and expected dispersal to decrease with acclimation period length. Post-release dispersal ranged from 2 to 90 km: ranging oryx acclimated for ca. 6 months moved 40-60 km from the release site, while penned oryx acclimated for ca. 1 month remained within 5-25 km. Our results demonstrate that captive management and environmental conditions at release strongly influence the extent to which reintroduced oryx disperse and adapt to wild conditions. We also show that-in contrast to previous studies-longer acclimation periods do not necessarily lead to site fidelity. Finally, our findings demonstrate the importance of tracking a large proportion of reintroduced individuals to (1) accurately record post-release behaviors and vital rates, and (2) adaptively evaluate pre-and post-release management actions to improve conservation outcomes.
Article
Full-text available
Bighorn sheep (Ovis canadensis) were once extirpated from the Black Hills region of South Dakota, USA, mirroring declining populations throughout North America. Since the 1960s, several reintroductions have occurred in the Black Hills to reestablish populations with varying success. We translocated 26 bighorn sheep from Alberta, Canada to the Black Hills (February 2015) to restore bighorn sheep to their historic range. Due to prior examinations of cause-specific survival, subsequent genetic diversity and disease prevalence were required to evaluate success of the restoration effort. We measured a mean allelic diversity of 5.23 (SE=0.44 [mean number of alleles]) and an observed heterozygosity of 0.71 (SE=0.06; expected = 0.64 ± 0.05) in the translocated individuals. Translocated bighorn sheep tested negative for Mycoplasma ovipneumoniae at capture. An autogenous vaccine was administered prior to release in an attempt to safeguard the translocated bighorn sheep from infection with a strain known to be resident in adjacent bighorn sheep populations. However, the year following the translocation, a different strain of M. ovipneumoniae was associated with a pneumonia outbreak that resulted in 57.9% mortality. Our results suggest that allelic diversity and heterozygosity were sufficient for long-term herd establishment, reducing the potential for founder effects. However, the This article is protected by copyright. All rights reserved. Deadwood bighorn sheep disease and diversity overwhelming mortality associated with pneumonia, via the transfer of M. ovipneumoniae from an unknown source, limited the success or our reintroduction efforts. Successful attempts to restore bighorn sheep to their historic ranges must consider and mitigate potential routes for M. ovipneumoniae transmission pre-and post-reintroduction.
Article
Full-text available
ABSTRACT—Bighorn Sheep (Ovis canadensis) historically inhabited the Black Hills region of South Dakota, but the species was extirpated from the area in the early 1900s concurrent with declines in population throughout their entire North American range. Translocation is a common management tool allowing for accelerated colonization of historic Bighorn Sheep habitat, but many attempts are unsuccessful. Mountain Lions (Puma concolor) and pneumonia are generally considered the most common limiting factors to Bighorn Sheep populations. Twenty-six Bighorn Sheep were translocated from Alberta, Canada to the Deadwood region of the northern Black Hills, an area with both a resident Mountain Lion population and potential for contact with Domestic Sheep (Ovis aries) and Goats (Capra hircus), known carriers of pathogens that are lethal to Bighorn Sheep. Adult survival and natality increased the population substantially in year 1, however, the only breedingage male was euthanized owing to concerns about potential for pathogen transmission from Domestic Sheep and Goats. In year 2, the population experienced a pneumonia outbreak, resulting in 57.9% of all mortalities during the study period. Mountain Lion predation was not detected, nor was direct contact with Domestic Sheep or Goats observed. Intensive monitoring was critical in determining the outcome of the translocation.
Article
Full-text available
ABSTRACT.—Initial movements of re- introduced wildlife populations can determine short-term restoration success. Managers need ways to encourage release site fidelity to mitigate suboptimal breeding, reduce mortality rates, and minimize human-wildlife conflicts. We studied initial movement ecology of elk (Cervus elaphus) fitted with GPS collars and re- introduced to the Missouri Ozarks in 2011 (n=32), 2012 (n=21), and 2013 (n=31) for 6 mo post release. We assessed maximum displacement from the release site, range shifts, and range size across four sequential time frames (0-10 d, 11-31 d, 32-61 d, and 62-183 d). Compared to other elk restorations in eastern North America, site fidelity was high, with maximum distance from the release site 62-183 d post release ≤10 km for 94% of 2011 animals, 57% of 2012 animals, and 97% of 2013 animals. Elk range sizes were similar during the first 61 d post release but doubled in size 62-183 d post release to an average of 26.2 km2 (range: 4.0—218.8 km2). The average range overlap for individual elk in sequential time periods was between 23-26% across years, indicating elk used different areas over time. Release site had the greatest influence on initial movements; one site used in 2012 was associated with greater release site displacement and range sizes. Maternal cows also demonstrated higher site fidelity to the release site than nonmaternal cows. High site fidelity and small home ranges in elk recently restored to Missouri may be attributed to soft release, minimal human disturbance, quality habitat, and release groups of mature females.
Article
Full-text available
Mountain goats (Oreamnos americanus) are among the least studied North American ungulates. Aided by successful translocations from the early to mid-1900s, introduced populations have greatly expanded within non-native ranges, yet there remains a paucity of empirical studies concerning their habitat requirements and potential distributions. The lack of studies presents a formidable challenge to managers tasked with monitoring mountain goat expansion and mitigating for any potential negative impacts posed to native species and communities. We constructed summer and winter resource selection models using GPS data collected during 2011–2014 from 18 (14 female and four male) mountain goats in the Snake River Range of the southwest Greater Yellowstone Area. We used generalized linear mixed models and evaluated landscape and environmental covariates at multiple spatial grains (i.e., neighborhood analyses within 30-, 100-, 500-, and 1000-m buffers) within four related suites. The multi-grain resource selection function greatly improved model fit, indicating that mountain goat resource selection was grain dependent in both seasons. In summer, mountain goats largely selected rugged and steep areas at high elevations and avoided high solar radiation, canopy cover, and time-integrated normalized difference vegetation index (NDVI). In winter, mountain goats selected lower elevations characterized by steep and rugged slopes on warm aspects and avoided areas with high canopy cover, NDVI amplitude, and snow water equivalent. Slope was the dominant predictor of habitat use in both seasons, although mountain goats selected for steeper slopes in winter than in summer. Regional extrapolations depicted suitable mountain goat habitat in the Snake River, Teton, Gros Ventre, Wyoming, and Salt Ranges centered around steep and rugged areas. Winter range was generally characterized by the steepest slopes within a more broadly distributed and generally less steep summer range. Further research should examine the spatial and temporal overlap with native populations to further our understanding of resource selection dynamics and the potential for introduced mountain goats to alter intraguild behavioral processes of sympatric species, namely the Rocky Mountain bighorn sheep (Ovis canadensis canadensis).
Book
The book covers population dynamics, diet, nutrition, diseases, behavior, and genetics of mountain lions occupying the Black Hills region. It explores the impact of a changing prey base on population growth and decline, movements within and away from the region, and hunting on the species; discusses interactions between the cats and livestock; and examines local people’s evolving perceptions of mountain lions. Provides a unique look into how a large, secretive predator recolonized an isolated region of North America.