Ecology and behavior of mule deer on the Rosebud Coal Mine, Montana
by Duane E Fritzen
A thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in
Biological Sciences
Montana State University
© Copyright by Duane E Fritzen (1995)
Abstract:
Mule deer (Odocoileus hemionus) inhabiting the Rosebud Coal Mine near Colstrip, Montana were
studied 1992-1995. Aerial surveys were used to assess distribution and abundance of deer.
Radiotelemetry provided information regarding deer movement, activity, use of vegetation-cover types,
and survivorship. Biological materials obtained from collected deer provided information regarding
food habits, physical condition, and productivity.
Based on aerial surveys, male and nonproductive female groups exhibited similar distributions different
from that of productive female groups. Deer population density, approximately 7.5 deer / km2 during
this study, increased 1974-1994, while deer distribution shifted from outlying portions of the study area
to core reclamation sites.
Fifty of 55 radiocollared deer monitored during this study were yearlong residents of the study area.
Mean home range size of males exceeded that of females annually and during all seasons. Diel mobility
and activity of females during summer, fall, and spring exceeded that during winter. Mobility and
activity were greatest during nocturnal, afternoon, and diurnal hours during summer, winter / fall, and
spring, respectively. Deer preferred pine savannah, riparian, and reclamation vegetation-cover types
and avoided mixed shrub and disturbance types. Annual survivorship of radiocollared fawns, adult
females, and adult males averaged 43.1, 90.0, and 57.7%, respectively. Leading causes of death for
fawns and adult females were coyote predation (58.6%) and vehicle collision (17.2%). Hunter harvest
accounted for 87.5% of all adult male deaths.
Examination of collected deer indicated forbs comprised 88.2; 50.5, and 55.9% of diets during summer,
fall, and spring, respectively. Browse predominated during winter, forming 79.4% of diets. Physical
condition of deer was greatest during fall and least during spring. Female ovulation and fertilization
rates were high, averaging 1.68 ova / female and 100.0%, respectively.
Although deer abundance increased since 1974, population characteristics during this study suggested a
relatively high-density but stable population. Deer movements suggested the study area provided
high-quality habitats capable of supporting deer on yearlong home ranges. Post-mining reclamation
was used extensively and consistently by deer. Collectively, these findings suggest that surface mining
on the study area benefited deer, at least in the short term, providing increased diversity of habitats
resulting in increased abundance of deer. ECOLOGY AND BEHAVIOR OF MULE DEER ON THE
ROSEBUD COAL MINE, MONTANA
by
Duane E Fritzen
A thesis submitted in partial fulfillment
of the requirements for the degree
of
Doctor of Philosophy
in
Biological Sciences
MONTANA STATE UNIVERSITY
Bozeman, Montana
December 1995
^ cIicI
APPROVAL
of a thesis submitted by
Duane E Fritzen
This thesis has been read by each member of the thesis committee and
has been found to be satisfactory regarding content, English usage, format,
citations, bibliographic style, and consistency, and is ready for submission to the
College of Graduate Studies.
Approved for the Major Department
Date
Head, Major Depgrfment
Approved for the College of Graduate Studies
Date
Graduate Dean
iii
STA TEM EN T OF PERMISSION TO USE
In presenting this thesis in partial fulfillment of the requirements for a
doctoral degree at Montana State University, I agree that the Library shall make it
available to borrowers under rules of the Library. I further agree that copying of
this thesis is allowable only for scholarly purposes, consistent with “fair use” as
prescribed in the U S. Copyright Law. Requests for extensive copying or
reproduction of this thesis should be referred to University Microfilms
International, 300 North Zeeb Road, Ann Arbor, Michigan 48106, to whom I have
granted “the exclusive right to reproduce and distribute my dissertation for sale in
and from microform or electronic format, along with the right to reproduce and
distribute my abstract in any format in whole or in part.”
Signature
Date
Z?. /993
ACKNOW LEDG M ENTS
The assistance of numerous individuals from Western Energy Co.,
Montana Power Co., Schwend Aviation, Montana State University, and the
Montana Department of Fish, Wildlife, and Parks is gratefully acknowledged.
Special thanks are extended to Richard Mackie, who provided guidance in all
aspects of the study, and Bill Schwarzkoph and Bruce Waage, who provided
technical support, assistance, and valuable advice throughout the study. Thanks
also go to graduate committee members Lynri Irby, Pat Munholland, Bret Olson,
Bob White, and Richard Horsweil for helpful comments regarding study design
and data collection procedures and for reviews of this thesis. The study was
funded by Western Energy Company, Colstrip, Montana.
vi
TABLE OF C O NTENTS
r
Page
APPRO VAL............'..............................................................
.................................
ii
STATEM ENT OF PERMISSION TO U S E .................................................
iii
VITA . . .............................................................................................................................
A C K N O W L E D G M E N T S ...................................
iv
v
TABLE OF C O N T E N T S ......................................................................
LIST OF T A B L E S .....................................................................................................
viii
LIST OF F IG U R E S ...............................................................................
A BSTRA C T..........................................................
xvi
IN TR O D U C TIO N .............................................................................................................
1
STU D Y AREA..................................................................................................................
4
Location.........................................................................................................
Regional and Local Geology......................................................................
Soils....................................................................................................
Climate and W e a th e r..................................................................................
Vegetation.....................................................................................................
W ildlife............................................................................................................
Land U s e ............................................................................................................
4
4
6
7
9
9
10
M E T H O D S ............
The Biological Y ear.................................................................................
Classification of Vegetation-Cover Types ....................................................
Deer C ap ture............................................................................
Radiotelemetry..................................................................................................
Aerial S urveys............................................................................................
Ground Surveys.............................................................
Deer Collections. ......................................................................................
Analytical Methods.................................................................................
12
12
14
16
19
20
vii
TABLE OF C O N TEN TS - Continued
Page
Deer Distribution Among Grid Cells ........................................
Deer Use of Vegetation-Cover Types..........................................
Home Range.......................................................................................
Abundance.........................................................................................
Survival................................................................................................
Statistical Comparisons........................
20
22
23
23
23
24
RESULTS & D ISCUSSIO N.......................................................................................
25
i
Vegetation-Cover T ypes..........................
Spatial Distribution of Deer.........................................................................
General Movement Patterns......................................................................
Home Range, Mobility, and Activity..................
Use of Vegetation-Cover Types by Radiocollared D e e r ......... >...........
Nocturnal Use of Colstrip...........................................................................
Food H abits..................................................................................................
Physical Condition.......................................................................................
Population Characteristics..........................................
Abundance and T re n d ....................................................................
Age / Sex Structure.........................................................................
Social Organization..............................................
Conception and Birth D a te s ...........................................................
Ovulation and Fertilization R a te s .................................................
Secondary Sex R a tio ......................................................................
Fawn Survivorship..............................
Adult Survivorship.............................................................................
MANAGEMENT IM P L IC A T IO N S ..............................................................................
25
35
52
54
59
74
79
82
83
83
87
89
90
92
92
94
94
100
Management in Relation to H ab itat............................................................
100
Management in Relation to H um ans...........................................................
103
Future Research N e e d s ................................................................................... 106
REFERENCES C ITE D .....................................................................
A PPEN D IX......................................................................................................................
126
viii
LIST O F TABLES
Table
1.
2.
3.
4.
5.
6.
7.
8.
9.
Page
Frequency of occurrence (%) of grasses among 100 1-m2
circular plots in each of 8 vegetation-cover types, Rosebud
Mine / Colstrip study area, July 1994...................................................
26
Frequency of occurrence (%) of forbs among 100 1-m2
circular plots in each of 8 vegetation-cover types, Rosebud
Mine / Colstrip study area, July 1994.................................................
28
Frequency of occurrence (%) of shrubs / trees among 100
100-m2 circular plots in each of 8 vegetation-cover types,
Rosebud Mine Z Colstrip study area, July 1 9 9 4 .................................
32
Proportional availability and use of grid cells containing
specific vegetation-cover types by mule deer (all age / sex
classes), Rosebud Mine / Colstrip, Montana, summer 1992spring 1994................................................................................ ............
40
Proportional availability and use of grid cells containing
specific vegetation-cover types by adult male mule deer, Rosebud
Mine / Colstrip, Montana, summer 1992-winter 1993.......................
42
Proportional availability and use of grid cells containing
specific vegetation-cover types by adult female mule deer without
fawns, Rosebud Mine / Colstrip, Montana, summer 1992winter 1993................................................................................................
45
Proportional availability and use of grid cells containing
specific vegetation-cover types by adult female mule deer with
fawns, Rosebud Mine I Colstrip, Montana, summer 1992winter 1993...................................................
47
Mean annual, seasonal, and diel home range sizes (ha)
for adult mule deer, Rosebud Mine / Colstrip, Montana,
fall 1992-spring 1994. . .........................................................................
55
Mean number of meters traveled per hour by adult female
mule deer during diel periods 1-4, Rosebud Mine / Colstrip,
Montana, summer 1992-spring 1994............................
57
ix
Table
1o:
11.
12.
13.
14.
15.
16.
17.
18.
19.
. Page
Mean percentage of radiocollared adult female mule deer active
during die! periods 1-4, Rosebud Mine / Colstrip1 Montana,
summer 1992-spring 1994......................................................................
58
Proportional occurrence of vegetation-cover types within home
ranges of radiocollared adult female mule deer in relation to
proportional availability of vegetation-cover types within the study
area, Rosebud Mine / Colstrip, Montana, summer 1993spring 1 9 9 4 ................................................................................................
60
Proportional occurrence of vegetation-cover types within home
ranges of radiocollared adult male mule deer in relation to
proportional availability of vegetation-cover types within the study
area, Rosebud Mine / Colstrip, Montana, summer 1993spring 1994..............................................................................
62
Proportional occupation of vegetation-cover types within home
ranges by adult female mule deer in relation to proportional
availability of vegetation-cover types within home ranges, Rosebud
Mine / Colstrip, Montana, summer 1993-spring 1 9 9 4 ........................
64
Proportional occupation of vegetation-cover types within home
ranges by adult male mule deer in relation to proportional
availability of vegetation-cover types within home ranges, Rosebud
Mine / Colstrip, Montana, summer 1993-spring 1994..........................
66
Number of radiolocations obtained from adult female mule deer
while occupying each of 8 vegetation-cover types during diurnal and
nocturnal periods, Rosebud Mine / Colstrip, summer 1993spring 1994..................................................................................................
70
Number of radiolocations obtained from adult female mule deer
while active and inactive in each of 8 vegetation-cover types,
Rosebud Mine / Colstrip, summer 1993-spring 1994............................
72
Mean Urban Deer Index (UDI) values for mule deer among
sections of Colstrip, Montana, July 1993-June 1994............................
75
Percentage occurrence of houses, mobile homes, and apartments
among sections of Colstrip, Montana, January 1994............................
76
Percentage occurrence of residences containing landscape
plantings, protected plantings, and deer-damaged plantings in
Colstrip, Montana, January 1994...............................................................
77
X
Table
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
Page
Mean percentage composition of diet items, by major forage
( class, and frequency occurrence (%) of individual plant
species contained within rumens obtained from female mule
deer, Rosebud Mine / Colstrip, Montana, fall 1992summer 1994................................................................................................
Means of whole and dressed weights and kidney fat index values
for adult female mule deer, Rosebud Mine / Colstrip, Montana,
fall 1992-summer 1994.........................
80
83
Lincoln index population estimates for mule deer, Rosebud Mine /
Colstrip, Montana, summer 1992-spring1995.........................................
84
Mule deer population density estimates (deer / km2) and relative
occurrence of fawns, adult females, and adult males derived
from aerial surveys (full-coverage helicopter flights only),
Rosebud Mine / Colstrip, Montana, winter 1993-spring 1995..............
88
Mean sizes and numbers of male, female, and mixed-sex
mule deer groups observed during ground surveys, Rosebud
Mine / Colstrip, Montana, June 1993-May 1994.......................................
91
Monthly survival estimates and 95% confidence intervals for
radiocollared female and male mule deer fawns, Rosebud Mine /
Colstrip, Montana, June 1992-May 1995.................................................
95
Seasonal causes of death for radiocollared female and male
mule deer fawns, Rosebud Mine / Colstrip, Montana, June 1993May 1 9 9 5 ..........................................
96
Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult female mule deer, Rosebud Mine /
Colstrip, Montana, June 1992-May 1995.................................................
97
Seasonal causes of death for radiocollared yearling / adult
female mule deer, Rosebud Mine / Colstrip, Montana, June
1992-May 1995................
97
Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult male mule deer, Rosebud Mine /
Colstrip, Montana, June 1992-May 1995...........................
98
xi
Table
30.
Page
Seasonal causes of death for radiocollared yearling / adult
male mule deer, Rosebud Mine / Colstrip, Montana, June 1992May 1995.........
99
/
31.
32.
33.
34.
35.
36.
37.
38.
39.
Capture / status of mule deer, Rosebud Mine / Colstrip,
Montana, March 1992-May 1995. . .........................................................
127
Monthly survival estimates and 95% confidence intervals for
radiocollared female and male mule deer fawns, Rosebud Mine /
Colstrip, Montana, June 1993-May 1994. .............................................
130
Monthly survival estimates and 95% confidence intervals for
radiocollared female and male mule deer fawns, Rosebud Mine /
Colstrip, Montana, June 1994-May 1 9 9 5 :..............................................
130
Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult female mule deer, Rosebud Mine /
Colstrip, Montana, June 1992-May 1993 .................................................
131
Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult female mule deer, Rosebud Mine /
Colstrip, Montana, June 1993-May 1994.................................................
131
Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult female mule deer, Rosebud Mine /
Colstrip, Montana, June 1994-May 1995................................
132
Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult male mule deer, Rosebud Mine /
Colstrip, Montana, June 1992-May 1993.................................................
132
Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult male mule deer, Rosebud Mine /
Colstrip, Montana, June 1993-May 1994.................................................
133
Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult male mule deer, Rosebud Mine /
Colstrip, Montana, June 1994-May 1995......... ...................................
133
z
/
xii
LIST OF FIGURES
Figure
1.
Page
The general region of study including mining areas, major
drainages and topographic features, and the city of C o ls trip ............
5
Monthly departures from the 30-year mean (1961-1990) with
respect to precipitation (cm) and temperature (C) patterns for
the period, June 1992-August 1994, Rosebud Mine (Colstrip
weather reporting station), Montana. The 30-year mean monthly
values, June-May, were: precipitation— June= 14.5, July=3.0,
August=S. 1, September=3.7, October=2.9, November=1.5,
December= 1.5, January=!?, February= 1.3, March=2.0, April=4.1,
and May=6.7; and temperature— June=17.8, July=217,
August=20.6, S e p te m b e r= # .! October=8.4, November=O.6,
December=-5.4, January=-G.8, February=-3.3, March= 1.2,
April=?.0, and May= 12.4.............................................................................
8
3.
Ground survey route through the Rosebud Mine, Montana................
18
4.
Sections A, B, C, D, and E of Colstrip in relation to Rosebud Mine
active mine / reclamation areas.................................................................
18
Distribution of mule deer groups on the Rosebud Mine / Colstrip
aerial survey area based on 8 fixed-wing aerial surveys (n=2 /
season), summer 1992 - spring 1 9 9 4 .......................
36
Summer distribution of mule deer groups on the Rosebud Mine /
Colstrip aerial survey area based on 2 fixed-wing aerial surveys
conducted during July 1992 and 1993......................................................
37
Winter distribution of mule deer groups on the Rosebud Mine /
Colstrip aerial survey area based on 2 fixed-wing aerial surveys
conducted during December 1992 and February 1993 .......................
37
Fall distribution of mule deer groups on the Rosebud Mine / Colstrip
aerial survey area based on 2 fixed-wing aerial surveys conducted
during October 1992 and 1993 ...............................................
............
38
2.
,
. 5.
6.
7.
8.
xiii
Figure
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
Page
Spring distribution of mule deer groups on the Rosebud Mine /
Colstrip aerial survey area based on 2 fixed-wing aerial surveys
conducted during March 1993 and May 1 9 9 4 .......................... .........
38
Mean number of mule deer observed per hour during aerial
surveys of the Rosebud Mine permit area, 1974-1994. Means
from 1974-1982 were derived from monthly surveys (n=12) flying
1-mile transects. Means from 1983-1985 were derived from
seasonal, full-coverage surveys (n=4). Means from 19861994 were derived from seasonal surveys (n=4) flying 0.5-mile
transects.........................................................................................................
86
Age-specific numbers of female and male mule deer obtained
from hunter harvest, deer-vehicle collision, and special collection,
Rosebud Mine / Colstrip, Montana, September 1992-June 1994. . . .
89
Monthly rates of survival for adult female and male mule deer,
Rosebud Mine / Colstrip, Montana, June 1992-May 1995 ...................
99
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer and winter 1974. . . . . .
134
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1975,
and spring 1976...............................
134
Distribution of mule deer groups on the Rosebud Mine permit area '
based on fixed-wing aerial surveys, summer, fall, and winter 1976,
and spring 1977..........................................................................................
135
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1977,
and spring 1978..........................................................................................
135
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1978,
and spring 1979. . ............ ....................... ' ..................................................
136
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1979,
and spring 1980.....................
136
xiv
Figure
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
Page
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1980,
and spring 1981...........................................................................................
137
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1981,
and spring 1982......................................................................................
137
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1982,
and spring 1983............................
138
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1983,
and spring 1984.........................
138
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1984,
and spring 1985..........................................................
139
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1985,
and spring 1986..............................................................................................
139
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1986,
and spring 1987....................................................................................
140
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1987,
and spring 1988..................................................................................
140
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1988,
and spring 1989..............................................................................................
141
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1989,
and spring 1990.......................
141
Distribution of mule deer groups on the Rosebud Mine' permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1990,
and spring 1991.....................
142
XV
Figure
30.
31.
32.
Page
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1991,
and spring 1992. ...............................................................................................
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1992,
and spring 1993..............................................................................................
Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1993,
and spring 1994........................................................
142
143
XVI
ABSTRACT
Mule deer fOdocoileus hemionus) inhabiting the Rosebud Coal Mine near
Colstrip, Montana were studied 1992-1995. Aerial surveys were used to assess
distribution and abundance of deer. Radiotelemetry provided information
regarding deer movement, activity, use of vegetation-cover types, and
survivorship. Biological materials obtained from collected deer provided
information regarding food habits, physical condition, and productivity.
Based on aerial surveys, male and nonproductive female groups exhibited
similar distributions different from that of productive female groups. Deer
population density, approximately 7.5 deer / km2 during this study, increased
1974-1994, while deer distribution shifted from outlying portions of the study area
to core reclamation sites.
Fifty of 55 radiocollared deer monitored during this study were yearlong
residents of the study area. Mean home range size of males exceeded that of
females annually and during all seasons. Diel mobility and activity of females
during summer, fall, and spring exceeded that during winter. Mobility and activity
were greatest during nocturnal; afternoon, and diurnal hours during summer,
winter / fall, and spring, respectively. Deer preferred pine savannah, riparian, and
reclamation vegetation-cover types and avoided mixed shrub and disturbance
types. Annual survivorship of radiocollared fawns, adult females, and adult males
averaged 43.1, 90.0, and 57.7%, respectively. Leading causes of death for
fawns and adult females were coyote predation (58.6%) and vehicle collision
(17.2%). Hunter harvest accounted for 87.5% of all adult male deaths.
Examination of collected deer indicated forbs comprised 88.2; 50.5, and
55.9% of diets during summer, fall, and spring, respectively. Browse
predominated during winter, forming 79.4% of diets. Physical condition of deer
was greatest during fall and least during spring. Female ovulation and fertilization
rates were high, averaging 1.68 ova / female and 100.0%, respectively.
Although deer abundance increased since 1974, population characteristics
during this study suggested a relatively high-density but stable population. Deer
movements suggested the study area provided high-quality habitats capable of
supporting deer on yearlong home ranges. Post-mining reclamation was used
extensively and consistently by deer. Collectively, these findings suggest that
surface mining on the study area benefited deer, at least in the short term,
providing increased diversity of habitats resulting in increased abundance of deer.
1
INTRODUCTION
Mule deer (Odocoileus hemionus') are widely distributed and extensively
studied in western North America. Investigations have been conducted in a wide
variety of natural environments including mountain-foothill (Robinette 1966, Ihsle
Pac et al. 1988, Pac et al. 1991), prairie (Dusek 1975, Severson and Carter
1978, Swenson et al. 1983, Wood 1986, Jackson 1990), breaks and badlands
(Mackie 1970, Komberec 1976, Dood 1978, Riley 1982, Kraft 1987, Hamlin and
Mackie 1989, Jensen 1992), and chaparral and desert (Linsdale and Tomich
1953, Kucera 1978, Bowyer 1984, Ordway and Krausman 1986) environments.
Studies also have been conducted in human-disturbed environments including
agricultural (Egan 1957, Ball 1987, O ’Connor 1987), industrial (Eberhardt et al.
1984, Clark and Medcraft 1986, Medcraft and Clark 1986) and urban (Mackie and
Pac 1980, Happe 1983, Vogel 1983, de Vos et al. 1984, Bellantoni et al. 1993)
environments. However, studies of mule deer occupying a complex of
undisturbed and highly disturbed environments generally are lacking.
The human population is expected to increase by 19.4% (3,709,000
individuals) in the Intermountain W est region of the United States between 1995
and 2010 (Bureau of the Census 1994). As human population increases,
development and alteration of natural habitats will continue such that mule deer
may be increasingly impacted (Reed 1981). Mule deer populations already have
decreased or been eliminated over portions of the species original distribution,
due primarily to recent human influences (Geist 1990). Accordingly,
2
understanding the ecology of mule deer inhabiting human-disturbed environments
is necessary if populations are to be maintained in the future.
The Rosebud Mine and town of Colstrip in southeastern Montana
collectively comprise an environment in which undisturbed native vegetation and
intensively disturbed agricultural, industrial, urban, and postmining reclamation .
sites are intermixed. Further, undisturbed and disturbed sites are highly
fragmented. The area is inhabited by a sizable population of Rocky Mountain .
mule deer (0 . h. hemionus), many members of which have access to the entire
range of sites along the disturbance continuum. Thus, the area represents a
unique environment in which to examine mule deer population-habitat
relationships in the presence of variable human disturbance.
This study was initiated in July 1991 by Western Energy Company
(W ECO), the owner and operator of the Rosebud Mine, in response to concerns
about mule deer encroachment upon industrial and residential sites on the
Rosebud Mine and in the town of Colstrip, respectively. The increased presence
of mule deer in these areas resulted in a greater incidence of deer-human,
interactions, many of which, including deer-vehicle collisions, deer damage to
gardens and ornamental shrubbery, and direct physical contact between deer and
humans, were detrimental to either humans or their property. The goal of the
study was to define habitat Z mule deer population relationships in this variable
environment. Specific objectives were to (1) evaluate characteristics of disturbed
and undisturbed habitats available to mule deer on the area, (2) define physical,
reproductive, survival, and behavioral characteristics of mule deer occupying the
3
area, and (3) assess mule deer distribution among and use of disturbed and
undisturbed vegetation-cover types on the area. Understanding populationhabitat relationships is necessary if W E C O is to achieve its long-term
management goals of (1) maintaining the mule deer population on the Rosebud
Mine, while (2) minimizing the occurrence of negative deer-human interactions in
Colstrip.
4
S TU D Y AREA
Location
The study was conducted primarily in the vicinity of the Rosebud Mine and
nearby town of Colstrip1 in southeastern Montana. Observations, however,
extended south to the Big Sky Mine and west to the Sarpy Creek Mine in Bighorn
County, yielding an overall study area of approximately 300 km2 (Figure 1). The
study area as defined specifically for analytical purposes was that area (130.2
km2) containing all relocations of radiocollared deer residing yearlong on or about
the Rosebud Mine. The Rosebud Mine lies at 45° 5 T north latitude and 106° 37'
west longitude and surrounds the town of Colstrip. The East Fork of Armell’s
Creek is the major drainage within the study area. It courses through the study
area along a predominantly east-west axis before turning north and eventually
emptying into the Yellowstone River, roughly 50 km north of Colstrip. Major
topographic features of the area include the Little Wolf Mountains and the Greenleaf Ridge which separate the Rosebud Mine from the Sarpy Creek and Big Sky
Mines, respectively.
Regional and Local Geology
The study area was located in the northern portion of the Powder River
Basin in the Northern Great Plains physiographic province (Plantenberg 1983).
Geologic history of the region since the Precambrian includes periods of
deposition, deformation, and erosion. Deposition occurred during the Paleozoic
5
Montana
State Highway 39 .
Kilometers
Colstrip
Rosebud Mine
Little Wolf Mountains'
divide
East Fork
Armell s Creek
Green-leaf Ridge / -
Big Sky Mine
Sarpy
Creek
Mine
Figure 1. The general region of study including mining areas, major drainages
and topographic features, and the city of Colstrip.
and Mesozoic when an epicontinental sea covered the area (Schafer et al. 1979).
The late Cretaceous marked the beginning of the Laramide orogeny, when the
Black Hills and Big Horn Mountains were uplifted and intermediate ground
subsided forming the Powder River Basin. Extensive deposition of sediments on
flood plains of the newly formed Powder River Basin during the Paleocene
resulted in development of the Fort Union Formation, a succession of rock strata
rich in coal deposits (Lewis and Roberts 1977). The Eocene was a period of
folding and faulting that produced many of the present structural features of
southeastern Montana (Glaze and Keller 1965). The region was buried by debris
from volcanic activity to the west during the Oligocene and Miocene. However,
6
uplift of areas bordering the Powder River Basin occurred again during the
Pliocene, rejuvenating streams throughout the basin and resulting in erosion that
continued through the Pleistocene, forming the present landscape. The northern
portion of the Powder River Basin was not glaciated during the Pleistocene, but
climate changes associated with glaciation doubtless contributed to increased
rates of erosion and greater relief of the landscape (Schafer et al. 1979).
The landscape in the vicinity of Colstrip (approximately 980 m elevation) is
characterized by rolling prairies with alternating ridges, drainages, and sandstone
bluffs (Meyn et al. 1976). Twenty to 50 percent of the land surface slopes less
than 8%, and relief ranges from 150-300 m (Schafer et al. 1979). Local geology
is dominated by the Tongue River member of the Fort Union Formation. The
Tongue River member is composed of lenticular sandstones, carbonaceous
shales, and porcelainite (scoria) beds overlying subbituminous coal seams
(Veseth and Montagne 1980). The coal seam underlying the Rosebud Mine is 58 m thick and lies under 5-65 m of sandy shale overburden (Dollhopf et al.
1977a).
Soils
The study area lies in a transition zone where Aridisols (desert soils) of the
southern Powder River Basin, Mollisols (prairie soils) of the eastern Great Plains,
and Alfisols (forest soils) of the Black Hills and Big Horn Mountains overlap.
Entisols (young, undeveloped soils) are common locally where steep slopes are
subject to erosion. In general, coarse-loamy and fine-loamy soil textural classes
7
predominate (Packer 1974).
Potential evapotranspiration far exceeds annual precipitation on the study
area, and run-off during snow melt and high-intensity thunderstorms prevents
some precipitation from ever entering the soil profile (Schafer et al. 1979).
Accordingly, soils are dry in the root zone during the majority of the growing
season. This lack of water penetration into the soil results in the accumulation of
lime .and other soluble salts near the base of the root zone (Schafer et al. 1979).
Because of these and other soil profile characteristics, Hertzog (1983) assigned
the study area to Land Capability Class IV, lands suitable for improved pasture
and rangeland but not suitable for row crops.
~
Climate and W eather
The study area has a semi-arid, continental climate characterized by
extreme temperatures and precipitation. January, and July typically are the
coldest and warmest months with long-term means of -6.8 and 21.7 C,
respectively (N.O.A.A. 1961-1990). Annual temperature extremes may range
between -40 and 40 C. February and June are the driest and wettest months
with long-term means of 1.3 and 14.5 cm, respectively. Annual precipitation may
range between 22 and 63 cm, but the long-term mean was 46 cm. Approximately
3A of the annual precipitation total falls during April-October and results mainly
from showers and high-intensity thunderstorms (Schafer et al. 1979). The
average frost-free season in southeastern Montana is approximately 110 days
(Wood et al. 1989), although most growth of native herbaceous plants occurs
8
during April-June (Smoliak 1956).
Seasonal weather during the 1992-1995 study period was characterized by
(1) warm, dry springs, (2) cool, dry summers, and (3) normal falls and winters
relative to long-term temperature and precipitation means (N.O.A.A. 1992-1995)
(Figure 2). Exceptions were (1) summer 1993, which was wetter than normal,
(2) fall 1993 and 1994, which were drier and wetter than normal, respectively, and
(3) winter 1992-1993, which was cooler than normal.
J J A S O N D J
F M A M J
J A S O N D J F M A M J
J A
June 1862 - August 1994
Precipitation (cm)
[ |
Temperature (C)
Figure 2. Monthly departures from the 30-year mean (1961-1990) with respect to
precipitation (cm) and temperature (C) patterns for the period, June 1992-August
1994, Rosebud Mine (Colstrip weather reporting station), Montana. The 30-year
mean monthly values, June-May, were: precipitation— June= 14.5, July=3.0,
August=3.1, S e p te m b e rs .7, October=2.9, November= 1.5, December= 1.5,
January=I.7, February=!.3, March=2.0, April=4.1, and May=6.7; and
temperature— June=17.8, July=21.7, August=20.6, September= 14.1,
O c to b e rs .4, N ovem b er0.6, D ecem b er-5.4, January=-6.8, February=-3.3,
March= 1.2, April=7.0, and May=12.4.
9
Vegetation
Native plant communities in southeastern Montana have been
characterized broadly as either mixed prairie or coniferous woodland, with
scattered intrusions of riparian vegetation on more mesic sites (Payne 1973,
Dollhopf et al. 1977b). Because large-scale manipulation of native vegetation
commonly occurs on surface mines, however, the Rosebud Mine / Colstrip study
area was a highly interspersed mosaic of native and introduced / human-created
vegetation types.
Wildlife
Mule deer were by far the most abundant of all big game species on the
study area, 1992-1995. White-tailed deer fOdocoileus virginianus) and elk
fCervus elaphus) were rare, and a lone moose briefly visited the area during
summer 1994. In addition to wild ungulates, W EC O maintained a captive
population of approximately 20 bison (Bison bison) on the Rosebud Mine. Upland
game birds were abundant and included sharp-tailed grouse (Tvmpanuchus
phasianellus). pheasant (Phasianus colchicus), and wild turkey (Meleagris
aallopavo). Sage grouse (Centrocercus urophasianus) and Hungarian partridge
(Perdix perdix) were observed occasionally. Coyotes (Canis latrans) were
abundant while bobcats (Felis rufus) and red fox (Vuloes vulpes) were sighted
infrequently. Large predators apparently were absent, although mountain lions
(Felis concolor) and black bears (Ursus americanus) were harvested in heavily-
10
wooded regions south and west of the study area.
Recreational hunting was a popular activity on public and private lands in
the Colstrip area. However, federal and state regulations and concern for the
safety of employees and equipment precluded hunting on the Rosebud Mine
except under limited, highly controlled conditions. Similar concerns generally
precluded recreational hunting in and around Colstrip, although limited bow
harvest of deer occurred within the townsite.
Land Use
Cattle grazing constitutes the primary historic land use of native vegetation
in the Colstrip area. Although surface coal mining was the major land use on the
Rosebud Mine, reclaimed lands also were subject to limited grazing. A small
amount of land adjacent to the mine was used by the town of Colstrip for family
dwellings, schools, commercial developments, parks, recreational developments,
and roads. Power-generating facilities, associated settling and cooling ponds,
pipelines, and service roads were located in and around Colstrip. Only a small
amount of native mixed prairie, both on and off the Rosebud Mine, was converted
to agricultural cropland, predominantly alfalfa (Medicago sativa).
Commercial development of coal on the study area began in 1917 when
Northern Pacific Railway Company (NPRC) developed a coal field to obtain highquality, low-cost coal to power its steam locomotives. To facilitate development,
a branch rail line was constructed to the coal field from the main line at Forsyth,
Montana in 1923. At that time, the coal field site became known as Colstrip.
11
Coal was first mined at Colstrip in 1924 by the Northwestern Improvement
Co., a subsidiary of NPRC. Production continued until 1958 when NPRC
switched from coal to diesel-powered locomotives. Mining resumed in 1968 when
W E C O 1 a subsidiary of Montana Power Co., purchased the operation, including
the town of Colstrip1 and began producing coal for electrical utilities in Montana
and several midwestern states (Schafer et al. 1979, Plantenberg 1983).
In 1971, Montana Power Co. and Puget Sound Power and Light Co.
constructed 2 350-megawatt coal-fired power-generating facilities in Colstrip
(Aasheim 1981). Two 700-megawatt facilities were added in the early 1980s
(Knudson and Peterman 1980). As a result of industrial development, residential
and commercial sections of Colstrip were developed, and the human population
increased to its present level of approximately 3,035 (U S. Bureau of the Census
1992). Production of coal from the Rosebud Mine also increased, peaking during
the early 1990s at roughly 12 million tons of coal extracted per year; 30% was
exported to Minnesota and Wisconsin and 70% was burned in Montana,
principally in Colstrip.
12
M ETHODS
The Biological Year
Seasonal designations were derived from changes in climatic conditions
and deer behavior. The biological year, 01 June-31 May, was divided into 4
seasons: (1) summer, 01 June-31 August; (2) fall, 01 September-30 November;
(3) winter, 01 December-28 February; and (4) spring, 01 March-31 May. All data
and analyses relating to seasonal patterns reflected these designations.
Classification of Vegetation-Cover Types
Eight vegetation-cover types were defined based on conspicuous
vegetation, topography, and dominant land-use characteristics. The amount of
the study area covered by each type was determined via the map-weight method
(White and Garrott 1990) following ground-truthing of aerial orthophoto maps.
Plant species composition was estimated for vegetation-cover types to
elucidate the unique vegetative characteristics of each. Frequency occurrence of
species was determined based on plots (n=100 / type) positioned randomly
throughout the study area (Gysel and Lyon 1980, Bonham 1989, Higgins et al.
1994). One- and 100-m2 circular plots were used to measure presence of
grasses and forbs and shrubs / trees, respectively. Circular plots were used to
reduce circumferenceiarea ratios, thereby reducing edge and the potential bias
introduced by researcher decisions to include or exclude individual plants from
plots (Bonham 1989). Determination of plot size was based on recommendations
13
of Bonham (1989) for grasses and forbs and Costing (1956) and Irwin and Peek
(1979) for shrubs / trees. Plots were examined during mid-summer (July 1994) to
maximize the probability of including both cool- and warm-season plant species.
Common and scientific names of plants followed Booth (1950) and Booth and
Wright (1962) except where otherwise noted.
Deer Capture
Sixty-seven yearlings, adults and fawns ^6 months of age (48 females and
19 males) were captured using cannon nets (Hawkins et al. 1968), drive nets
(Beasom et al. 1980), a dart gun (Pond and O Gara 1994), and a hand-held net
gun (Barrett et al. 1982), March 1992-January 1993 (Appendix, Table 1).
Captured deer were sexed, aged according to tooth replacement and wear
criteria (Robinette et al. 1957, Rees et al. 1966), marked with metal ear tags, and
fitted with uniquely colored neck bands equipped with radiotransmitters (Telonics,
Inc., Mesa, Ariz.) (n=58) or neck bands / plastic ear flags without
radiotransmitters (n=9) (Nietfeld et al. 1994).
Forty-one neonate fawns (11 females and 30 males) were captured using
either random ground searches and behavioral cues of adult females (Downing
and McGinnes 1969, White et al. 1972, Huegel et al. 1985) or aerial location
techniques (Riley and Dood 1984), June 1993 and 1994 (Appendix, Table 1).
Captured fawns were sexed, weighed, marked with metal ear tags, and fitted with
either expandable, break-away neck bands equipped with radiotransmitters
(n=34) or uniquely marked plastic ear flags without radiotransmitters (n=7)
14
(Nietfeld et al. 1994).
Radiotelemetrv
Use of vegetation-cover types and patterns of home range establishment,
mobility, activity, and survival were determined through analysis of radiotelemetry
data. Ninety-two radiocollared deer were located 9,821 times through aerial
telemetry (n=151 locations, 1.5%) and ground triangulation (n=9,670 locations,
98.5%). Aerial locations were obtained using a Piper Supercub fixed-wing aircraft
equipped with 2-element H-antennas mounted on each wing strut. Deer location
was determined by homing in on the transmitter signal and obtaining visual
verification of the position of the radiocollared deer (Gilmer et al. 1981, Mech
1983, Kenward 1987, Samuel and Fuller 1994). The error polygon method
(Heezen and Tester 1967, Nams and Boutin 1991) was used to estimate deer
location from the ground using a single hand-held, 2-element H-antenna. A
Telonics TR-2 receiver was used for both aerial and ground telemetry (Telonics,
Inc., Mesa, Ariz.). All locations were recorded to the nearest 10 m as Universal
Transverse Mercator (UTM) grid coordinates.
Deer activity status (active or inactive), recorded at the time of location,
was determined via visual verification or estimated based on variability of
radiosignal strength (Kjos and Cochran 1970, Gilmer et al. 1971, Cochran 1980).
Fluctuating radiosignal strength indicated deer activity whereas constant signal
strength indicated inactivity.
Telemetry error was evaluated throughout the study by visually verifying
15
the location and activity status of randomly selected radiocollared deer
subsequent to remote telemetry contact and estimation of location and activity
(Samuel and Fuller 1994). Error trials were conducted during both diurnal and
nocturnal hours and during each season.
Mean error of telemetry bearings was
2.2±2.6 (SE) degrees based on 76 error trials conducted at a mean receivertransmitter distance of 376.8 m. This error resulted in misclassification of
vegetation-cover type occupied by deer 2.6% of the time. In contrast to the
findings of Gillingham and Bunnell (1985) and Beier and McCullough (1988),
activity status of deer based on fluctuating radiosignal strength was reliably
estimated (94.5% correct estimation based on 127 error trials).
Radiocollared deer were located 4 times monthly to assess seasonal
patterns of survival and home range use. One location per deer per month was
obtained during each of 4 different diel periods: (period 1) sunrise to mid day;
(period 2) mid day to sunset; (period 3) sunset to mid night; and (period 4) mid
night to sunrise. Successive locations obtained for each radiocollared deer were
separated by a minimum of 24 hours to ensure independence of observations.
A subset of radiocollared deer (n=16 yearling and adult females) was
monitored during diel periods to assess diel home range, mobility, and activity
characteristics among seasons. Diel monitoring involved location of deer at
hourly intervals spanning a 24-hour period. Attempts were made to monitor each
deer during at least 1 diel period per month. Deer monitored over diel periods
generally occupied home ranges on the periphery of Colstrip, centered near
active mining and reclamation sites. Accordingly, they had access to a wide
16
variety of vegetation-cover types and were accessible year-round.
Aerial Surveys
Deer population characteristics and distribution patterns were assessed
via counts and classifications of deer groups observed during aerial surveys of
the 23,550-ha Rosebud Mine permit area. Surveys were conducted seasonally
throughout the study using a Piper Supercub fixed-wing aircraft flown along northsouth transects spaced at half-mile intervals. Flights began at sunrise and were
terminated approximately 1.5 hours thereafter, or as weather dictated. Completecoverage aerial surveys (Hamlin and Mackie 1989) using either a Bell B-1 or
Hughes 500-D helicopter were conducted during winter 1993 and 1994 and
spring 1994 and 1995 to supplement data obtained during fixed-wing surveys.
Helicopter flights were initiated at sunrise and generally continued throughout the
day until sunset, with the exception of pauses for refueling. Deer groups
observed during flights were classified as to size, age / sex composition
(attempted during summer, fall, and winter surveys only), and vegetation-cover
type occupied. Location of groups was recorded as occurring in 1 of 910 25.9-ha
grid cells (804.5 x 321.8 m, 10 cells / mile2) comprising the survey area (Porter
and Church 1987, Hamlin and Mackie 1989, Jackson 1990). Deer age classes
were fawn (<12 months of age) and adult (;> 12 months of age). Identification of
individually marked radiocollared deer observed during flights was recorded.
Data from previous aerial surveys of the Rosebud Mine permit area,
including the number of deer observed per hour flight time and location of
17
observed deer groups, were obtained from W E C O Annual Wildlife Reports
(1974-1992) and used to estimate trends in mule deer population size and
distribution. Both transect and full-coverage surveys were conducted by W EC O
personnel using a Piper Supercub fixed-wing aircraft. Deer observed during
these flights were recorded by number and general location on a grid map of the
area.
Ground Surveys
Ground surveys were conducted along 2 independent vehicle routes
established on the Rosebud Mine and in the city of Colstrip to evaluate seasonal
patterns of social organization and deer use of urban areas, respectively.
Location, size, and age / sex composition when possible were recorded for all
deer groups observed during surveys. Four surveys were conducted monthly on
the Rosebud mine, 2 each during morning and evening crepuscular hours. The
survey route spanned 38.6 km and extended along an east-west axis through the
western two-thirds of the mine (Figure 3). Four nocturnal spotlight surveys were
conducted in Colstrip each month. Surveys were initiated between OOOO and
0100 hours M S T. The survey route included all major and selected side streets
in each of 5 sections of Colstrip (Figure 4); sections were defined according to
geographic position and dominant housing / landscape features. To compensate
for different lengths of the survey route among sections of Colstrip, data were
recorded as the number of deer observed per kilometer of survey route
(hereafter, Urban Deer Index [UDI]).
18
N
—Rosebud Mine Permit Area
—City of Colstrip
Kilometers
-Highway 39
--Survey Route
Figure 3. Ground survey route through the Rosebud Mine, Montana.
N o rth
M e te rs
S ta te H ig h w a y 3 9 ------
R o s e b u d M in e :
a c tiv e m in e &
r e c la m a tio n a r e a s
C a s tle
Rock
N Lake
Pow er
P la n ts
R o s e b u d M in e :
a c tiv e m in e &
r e c la m a tio n a r e a s
Figure 4. Sections A, B, C, D, and E of Colstrip in relation to Rosebud Mine
active mine / reclamation areas.
19
Characteristics of residences were determined for each section of Colstrip
to assess their potential impacts on spatial and temporal distributions of deer.
Four hundred residences were selected randomly from the city telephone
directory. Upon selection, residences were located and inspected from the street.
Information recorded included (1) type of residence, (2) presence / absence of
landscape plantings, (3) presence / absence of any device protecting plantings
from deer browsing, and (4) presence / absence of obvious deer damage to
plantings. Because residences were examined from front, the occurrence of
landscape plantings and deer damage likely was underestimated. The survey
was conducted during January 1994.
Deer Collections
Biological materials and physical measurements were obtained from 133
deer harvested by hunters (n=68), killed in deer-vehicle collisions (n=17), and
collected for study (n=48, 2-4 females collected per month). W hen carcass
condition allowed, materials collected included: (1) the lower jaw to determine age
using tooth replacement and wear criteria (Robinette et al. 1957, Rees 1966); (2)
kidneys and attached perirenal fat to calculate a kidney-fat index (KFI) (Riney
1955); (3) female reproductive tracts to determine conception dates (Hudson and
Browman 1959) and assess incidence of ovulation and fertilization based on
ovarian structures (Cheatum 1949); and (4) rumen contents for analysis of food
habits.
Food habits were estimated via analysis of 1-quart samples of rumen
20
contents obtained from hunter-harvested (n=39), vehicle-killed (n=5), and specialcollection (n=46) deer. Samples were fixed with 10% formalin and later rinsed,
separated, and examined macroscopicaliy at the Montana Department of Fish,
Wildlife & Parks Wildlife Research Lab in Bozeman, Montana. Rumen contents
were identified to the species level when possible and / or assigned to 1 of 3
major forage classes: (1) grasses; (2) fofbs; and (3) shrubs / trees. The pointframe method (Chamrad and Box 1964) was used to estimate the proportion of
each rumen sample comprised by each of the major forage classes.
Analytical Methods
Deer Distribution Among Grid Cells
Use versus availability analyses (Neu et al. 1974 and Byers et al. 1984)
were used to examine spatial distribution of deer groups observed during fixedwing aerial surveys in relation to the composition of vegetation-cover types within
occupied grid cells. Types contained within each grid cell were determined via
visual inspection of aerial photos. Expected use was defined as the proportion of
grid cells comprising the study area that contained the vegetation-cover type of
interest. Observed use was defined as the proportion of grid cells occupied by
deer groups of a given sex / productivity class that contained the type of interest.
Overall selection of grid cells was determined by summing chi-square goodnessof-fit test values over each vegetation-cover type. Selection of grid cells
containing specific types was determined by constructing 95% Bonferroni
confidence intervals about observed use proportions and noting the relative
21
position of expected use proportions.
Distributions of male and female deer groups in relation to the diversity of
vegetation-cover types contained within grid cells also was examined. The
number of vegetation-cover types contained within each grid cell was determined
via visual inspection of aerial photos, as above. Mean numbers of types
contained within grid cells occupied and not occupied by males and females were
compared using Wilcoxon rank sum tests. Mean numbers of types contained
within grid cells occupied by males and females were compared among seasons
using Kruskal-Wallis tests.
Spatial distribution of male, non-productive female, and productive female
deer groups among grid cells was examined seasonally using chi-square
goodness-of-fit tests. For each comparison, the expected value was the lesser of
the 2 numbers of grid cells occupied by each sex / productivity class being
examined. The observed value was the number of grid cells actually shared by
the 2 classes. For example, if male and productive female groups were observed
in 25 and 50 grid cells, respectively, during a seasonal aerial survey, they would
co-occur in 25 cells if the 2 classes exhibited complete spatial overlap and 0 cells
if they exhibited complete segregation. In this example, 25, the lesser of the 2
numbers of grid cells occupied by each class being compared, represents the
expected test value. Chi-square values were used to infer either aggregation /
segregation (significant values) or the lack thereof (non-significant values).
22
Deer Use of Vegetation-Cover Types
Use of vegetation-cover types by radiocollared deer was examined at both
second-order (placement of a home range within the study area) and third-order
(use of types within a home range) levels (Johnson 1980). Second-order
analyses (annual and seasonal) compared the proportion pf each vegetationcover type within the collective home range of all radiocollared deer (observed
use) to the proportion of each type comprising the study area (expected use).
For these analyses, the study area was defined as the area (12,016.4 ha) within
the perimeter of all annual home ranges of radiocollared deer residing yearlong in
the vicinity of the Rosebud Mine and city of Colstrip. Analyses followed the use.
versus availability procedures of Neu et al. (1974) and Byers et al. (1984). Chisquare goodness-of-fit tests determined overall selection or the lack thereof.
Bonferroni 95% confidence intervals then were constructed about observed use
proportions and compared to corresponding expected use proportions to
determine preference / avoidance of each vegetation-cover type.
Third-order analyses (seasonal only) compared the proportion of locations
for all radiocollared deer recorded within each vegetation-cover type (observed
use) to the proportion of each type comprising the collective home range of all
radiocollared deer with access to the type of interest (expected use). Because
radiocollared deer did not have equal access to all vegetation-cover types, chisquare goodness-of-fit-tests could not be used to determine overall selection.
Preference or avoidance, however, was determined by constructing a 95%
confidence interval about the observed use proportion for each vegetation-cover
23
type and comparing it to the corresponding expected use proportion.
Home Range
Annual, seasonal, and diel home ranges were calculated as minimum
convex polygons (Mohr 1947) using the HOME RANGE computer software
package (Ackerman et al. 1989). Annual, seasonal, and diel home ranges were
derived from 48, 12, and 25 radiolocations per deer, respectively.
Abundance
Deer numbers on the Rosebud Mine permit area were estimated via the
Lincoln index (Davis and Winstead 1980) using data obtained during both fixedwing and helicopter aerial surveys. The number of radiocollared deer in the
population available for sighting during each survey was determined either shortly
before or after each flight. Confidence limits calculated for abundance estimates
were derived according to Bailey (1951).
Survival
Seasonal and annual survival rates of deer were calculated using the
Kaplan-Meier or product-limit estimator (Kaplan and Meier 1958), modified to
allow staggered entry of radiocollared deer (Pollock et al. 1989). To eliminate
potential deaths from capture-related injuries or capture myopathy (Chalmers and
Barrett 1982, Conner et al. 1987), radiocollared adults, yearlings, and fawns >6
months of age were not considered at risk of mortality until 14 days post-capture.
The log-rank test (Cox and Oakes 1984, Pollock et al. 1989) was used to
24
compare survivorship functions of deer cohorts.
V
Statistical Comparisons
Statistical procedures were selected based on (1) the hypothesis tested
and (2) how well data being compared met the assumptions of each procedure.
Kruskal-Wallis nonparametric tests using chi-square approximations were used to
compare more than 2 data sets. Multiple comparisons were made using Dunn’s
test. T-tests and WiIcoxOn rank sum tests were used to compare 2 independent
data sets. Wilcoxon signed rank tests were used to make paired comparisons
between 2 dependent data sets. Chi-square goodness-of-fit tests were used to
analyze categorical data. Z-tests were used to compare binomial proportions. All
statistical analyses were accomplished using the Statistical Analysis System
(SAS Inst. Inc. 1987).
25
RESULTS & DISCUSSION
Vegetation-Cover Types
Vegetation of the Rosebud Mine / Colstrip area has been classified broadly
as ponderosa pine savannah featuring small islands of ponderosa pine (Pinus
ponderosal amidst expansive grasslands (Payne 1973). The 8 vegetation-cover
types defined for this study included 4 native (grassland, mixed shrub, pine
savannah, and riparian) and 4 human-created (disturbance, reclamation,
agriculture, and urban). The percentage occurrence of grass, forb, and shrub /
tree species varied markedly among types (Tables 1-3). In general, however,
diversity of plant species was highest in mixed shrub and riparian types, followed
by reclamation, pine savannah, grassland, disturbance, urban, and agriculture
types.
Grassland (7.4% of the study area) comparable to the central grassland
vegetation type described by Payne (1973) occurred on uplands with flat to gently
rolling topography. Native grasses, especially junegrass (Koeleria cristata) and
green needlegrass fStipa yiridula), dominated. Forb abundance and diversity
was intermediate relative to other vegetation-cover types. Shrubs were few and
widely dispersed.
Mixed shrub (31.1 %) was a combination of the Artemisia tridentata /
Aaropvron smithii and Artemisia cana / Aaropvron smithii vegetation types
described by Hansen et al. (1984). Silver sagebrush (Artemisia canal and big
sagebrush (A. tridentata) dominated structurally on both lowland and upland
Species
O
O
O
O
O
22
O
O
O
O
O
O
O
5
11
10
5
O
O
O
94
O
O
O
O
12
MX
O
1
O
O
. O
4
O
O
1
3
8
O
16
19
O
14
62
O
O
O
65
O
1
O
O
21
PS
O
O
O
O
O
O
.0
O
O
9
O
O
20
O
O
5
32
< O
O
O
21
O
O
O
O
11
RP
2
7
O
2
8
10
O
3
O
O
O
P
3
O
13
5
8
4
2
8
5
O
O
2
8
60
DS
O
3
O
O
3
5
O
O
O
O
O
O
O
O
2
13
10
O
O
16
4
O
15
O
O
3
RC
AG
O
1
3
14
O
34
4
3
1
O
O
1
2
O
39
. 15
O
O
O
O
O
5
O
O
O
O
O
6
O
O
O
28
17
O
_ O
O
O
O
O
O
O
18
.
6
O
O
O
1 .
2
5
O
O.
21
UR
0
0
0
0
0
3
0
0
0
0
0
0
15
13
moo- ' —c o o o o o
Aeailoos cylindrica
Aaropvron cristatum
Aaropvron dasvstachvum
Aaropvron elonaotum
Aaropvron repens
Aaropvron smithii
Aaroovron spicatum
Aaropvron trachvcaulum
Andropoaon aerardii
Andropoaon scoparius
Aristada Ionaiseta
Avena sativa
Bouteloua curtipendula
Bouteloua aracilis
Bromus inermis
Bromus japonicus
Bromus tectorum
Carex spp.
Echinochloa crusaalli
Hordeum jubatum
Koeleria cristata
Muhlenberaia cuspidate
Orvsoosis hvmenoides
Panicum capillare
Phalaris arundinacea
Poa pratensis
CD
Table 1. Frequency of occurrence (%) of grasses among 100 1-m2 circular plots in each of 8 vegetation-cover types,
Rosebud Mine / Colstrip study area, July 1994.
70
Table 1. Continued, 2 of 2.
Species
RP
O
4
41
7
O
2 .
O
35
32
2
DS
RC
AG
O
O
8
5
O
O
O
9
16
O
43
UR
O
PS
O
O
O
71 .
42
O
'
O
O
O
23
55
O
MX
O
Scirpus americanus
Sporobolus crvotandrus
Stipa comata
Stipa viridula
Triticum aestivum
GLa
0
a Designation of vegetation-cover types: GL=grassland, MX=mixed shrub, PS=pine savannah, RP=riparian,
DS=disturbance, RC=reclamation, AG=agriculture, and UR=urban.
M
-■ j
Table 2. Frequency of occurrence (%) of forbs among 100 1-m2 circular plots in each of 8 vegetation-cover types,
Rosebud Mine / Colstrip study area, July 1994.
Species
Achillea millefolium
Alvssum desertorum
Ambrosia psiIostachva
Antennaria microphvlla
Artemisia dracunculus
Artemisia friaida
Artemisia Iudoviciana
Asclepias pumila
Asclepias speciosa
Astraaalus adsuraens
Astraaalus americanus
Astraaalus barrii
Astraaalus bisulcatus
Astraaalus crassicarpus
Astraaalus drummondii
Astraaalus aracilis
Brassica spp.
Calochortus nuttallii
Camelina microcaroa
Camoanula rotundifolia
Cathartolinum riaidum
Centaurea maculosa
Ceratoides Ianata
Chenopodium album
Chrvsopsis villosa
Cirsium arvense
GLa
11
O
48
O
O
365
O
O
O
O
O
4
5
O
O
7
O
O
O
O
O
O
O
O
9
MX
31
12
15
2
13
46
3
2
O
9
O
3
O
2
1 ■
1
3
9
3 .
1
19
O
6
O
O
O
PS
RP
DS
RC
20
O
13
O
O
27
8
O
O
O
O
O
O
O
4
O
O
O
O
5
O
O
O
O
O
O
22
O
6
O
1
7
25
O
1
2
O
O
O
O
O
O
5
O
O
O
1
O
O
3
O
. 23
O
O
3
O
3
2
O
4
O
2
O
O
1
O
O
O
2
O
O
O
O
O
O
3
2
O
7
2
26
O
9
11
4
O
O
O
1
O
10
O
O
O
3
O
1
O
O
1
O
O
2
O
AG
4
O
21
O
O
O
O
O
O
O
O
O
O
O
O
O
8
O
O
O
O
O
O
O
O
O
UR
O
O
24
O
O
16
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
Table 2. Continued, 2 of 4.
Species
GLa
Cirsium undulatum
Convolvulus arvensis
Crvptantha celosioides
Descurainia sophia
Echinacea pallida
Erioeron spp.
Eriooonum flavum
Eriooonum pauciflorum
Ervsimum asperum
Euphorbia esula
Gaillardia aristata
Gaura coccinea
Glvcvrrhiza Iepidota
Grindelia squarrosa
Gutierrezia sarothrae
Hedvsarum boreale
Helianthus petiolaris "
Iva xanthifolia
Kochia scoparia
Lactuca pulchella
Lactuca serriola
Linum perenne
Lvoodesmia iuncea
Medicaoo Iupulina
Medicaoo sativa
Melilotus alba
Melilotus officinalis
3
O
O
O
15
O
4
O
7
O
O
3
O
O
O
' O
6
O
O
O
O
10
O
O
10
14
61
MX
10
O
3
O
20
6
O
2
4
1
1
1
1
4
20
6
O
O
O
3
11
8
1
O
2
1
11
PS
11
O
O
O
5
O
O
O
O
O
O
O
O
4
22
O
O
O
O
O
16
5
O
O
O
O
9
RF
2
5
O
1
5
O
O
O
O
O
O
O
O
1
3
O
O
6
O
1
21
1
O
2
18
3
13
DS
5
7
O
O
8
O
O
O
O
O
O
2
O
3
O
O
O
O
8
O
10
0
0
0
0
5
23
RC
1
14
0
0
2
0
0
0
1
0
0
5
0
5
0
3
1
0
0
0
4
35
0
1
21
3
51
AG
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0 -/
0
0
5
0
0
7
0
0
54
0
13
UR
0
13
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7
, 0
0
19
0
0
12
8
0
15
Table 2. Continued, 3 of 4.
Species
Mentzelia decapetela
Mertensia oblonaifolia
Monarda fistulosa
Oenothera serrulata
Oountia oolvacantha
Oreocarva sericea
Orthocarpus Iuteus
Oxvtropis spp.
Perideridia aairdneri
Petalostemon candidum
Petalostemon purpureum
Phlox hoodii
Plantaao pataaonica
Polyaala alba
Psoralea arqophylla
Psoralea esculenta
Psoralea tenuiflora
Ratibida columnifera
Rumex crispus
Salsola kali
Senecio plattensis
Solidaao missouriensis
Solidaao riaida
Sohaeralcea coccinea
Taraxacum officinale
Thlaspi arvense
Toxicodendron rvdberaii
GLa
MX
PS
RP
O
O
O
O
8
O
O
O
O
O
17
O
26
O
14
5
12
63
O
O
O
O
O
6
O
O
O
O
1
O
11
2
2
6
3
3
3
8
4
36
O
33
9
4
17
O
O
5
6
2
12
O
O
O
O
O
O
O
. O
O
O
18
O
6
25
O
4
O
41
6
O
13
O
O
O
5
9
O
O
O
O
O
O
7
3
O
O
1
5
O
O
O
2
O
O
6
2
1
8
15
O
O
O
7
O
7
11
2
.
.
DS
O
O
O
. O
.0
O
O
O
O
O
O
O
O
O
2
O
O
3
O
68
O
7
O
O
5
O
O
RC
1
O
O
O
O
O
O
1
O
1
1
O
1
2
8
O
1
13
O
6
O
1
O
O
4
1
O
AG
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
O
8
O
O
.O
O
3
O
O
UR
O
O
O
O
O
O
O
O
O
O
4
O
O
O
9
O
O
3
O
O
O
O
O
O
28
O
O
Table 2. Continued, 4 of 4.
Species
Yucca alauca
Traoopoaon dubius
Trifolium spp.
Trialochin spp.
Tvpha Iatifolia
Verbena bracteata
Urtica dioica
GLa
MX
PS
O
65
O
O
O
O
O
O
57
O
1
O
1
O
14
30
O
9
O
5
O
RP
O
7
O
O
11
O
8 .
DS
RC
O
12
O
O
O
O
O
O
34
O
O
O
1
O
AG
O
10
O
O
O
O
O
a Designation of vegetation-cover types: GL=grassland; MX=mixed Shrub; PS=pine savannah; RP=riparian;
DS=disturbance; RC=reclamation; AG=agriculture; and UR=urban.
UR
O
11
23
O
5
O
O
Table 3 Frequency of occurrence (%) of shrubs / trees among 100 100-m2 circular plots in each of 8 vegetation-cover
types, Rosebud Mine / Colstrip study area, July 1994.
Species
Artemisia cana
Artemisia tridentata
Atriplex canescens
Chrvsothamnus nauseosus
Chrvsothamnus viscid iflora
Eleaanus anaustifolia
Fraxinus pennsvlvanica
Juniperus Scopulorum
Juniperus horizontalis
Malus alaucescens
Pinus ponderosa
Populus deltoides ^
Prunus americana
Prunus virainiana
Rhus tri Iobata
Ribes spp.
Rosa spp.
Salix spp.
Shepherdia araentea
Svirnohoricarpos occidental is
Tamafix pentandra
GLa
MX
10
O
O
O
O
O
O
O
O
O
O
O
O
O
6
O
24
O
O
O
O
57
41
O
5
O
O
O
5
1
O
9
O
O
O
64
O
5
O
O
7
O
PS
RP
23
O
O
11
O
5
O
69
O
O
100
O
O
O
82
O
22
O
O
33
O
28
3
O
6
O
O
18 .
11
. O
O
2
18
7
33
28
8
35
18
1
50
2
DS
RC
AG
0
0
0
4
9
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
14
3
52
29
6
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
a Designation of vegetation-cover types: GL=grassland, MX=mixed shrub, PS=pine savannah, RP=riparian,
DS=disturbance, RC=reclamation, AG=agriculture, and UR=urban.
UR
10
0 .
0
0
0 .
19
0
23
0 Co
17 1x3
26
0
0
0
12
0
0
11
0
5
0
33
sites. Fragrant sumac (Rhus aromatica) (McGregor 1986) also occurred on
upland sites, especially those with rocky and / or less developed soils. Native
grasses and forbs were abundant and diverse.
Pine savannah (16.7%) occurred primarily on upland sites, particularly
those with rough topography and rocky soils (Payne 1973). Ponderosa pine,
Rocky Mountain juniper Uuniperus scopulorumV and fragrant sumac were
principal over and midstory species. Understory grasses, including native
bunchgrasses and introduced downy chess brome (Bromus tectorum), and forbs
were of intermediate abundance and diversity.
1
Riparian (3.1%) included all 4 formation classes (forest / woodland, shrub /
scrubland, herbland, and altered environments) described by Batchelor et al.
(1982) for riparian environments in Montana. Riparian vegetation was distributed
widely throughout the study area, occurring in lowland and upland mesic
drainages and sub-irrigated sites. Trees and shrubs such as green ash (Fraxinus
pennsvIvanicaT chokecherry (Prunus viminianaT wild plum (Prunus americana).
and common snowberry fSymphoricarpos albusl typically dominated. Understory
grasses and forbs were abundant and diverse and included roughly equal
numbers of native and introduced species.
Disturbance (19.1%), land disturbed by mining but not yet reclaimed,
included both raw spoil removed from open mining pits and sites from which
topsoil was removed to expose underlying coal. Disturbed sites were
characterized by a general lack of vegetative ground cover. Plant communities
that did establish often lacked diversity, containing primarily introduced, annual
34
grasses, including brome grasses CBromus spp.). and annual and biennial forbs
such as kochia CKochia scoparia). Russian thistle CSaIsoIa kali), prickly lettuce
CLactuca serriola) and yellow sweetclover CMeIiIotus officinalis). Shrubs were
rare, though rabbitbrushes CChrvsothamnus spp.) occurred on older sites.
Reclamation (18.1%) was characterized by high site to site variability in
plant species composition due principally to differences in seeding and
management practices (Lovell 1992). Reclamation was of 3 general types: (1)
grassland [85.0%]; (2) shrub grassland [14.7%]; and (3) pine grassland [0.3%].
Reclaimed grasslands were dominated by native and introduced grasses
including western wheatgrass CAaropvron smithii) (Sutherland 1986), tall
wheatgrass CAaropvron elonaatum) (Sutherland 1986), Kentucky bluegrass (Poa
pratensis). and smooth brome CBromus inermis). These grasses were present on
reclaimed shrub-grasslands as well, but structural diversity was enhanced by
fourwing saltbush CAtripIex canescens) and, to a lesser degree, rose (Rosa spp.)
and silver sagebrush. Pine grassland was similar vegetatively to shrub grassland
reclamation except ponderosa pine replaced fourwing saltbush as the major
structural component. Forbs were abundant and diverse, especially on reclaimed
grasslands, and consisted primarily of species found among native grassland and
mixed shrub vegetation-cover types.
Agriculture (2.7%) included fields of alfalfa CMedicaao sativa). wheat
(Triticum aestivum) (Sutherland 1986), and oats CAvena sativa) (Sutherland
1986) generally less than 15 ha in size and scattered throughout the study area.
Grasses and forbs commonly encountered among crops included brome grasses,
35
Kentucky bluegrass, yarrow (Achillea millefolium), western ragweed (Ambrosia
psilostachva). mustard (Brassica spp.) and yellow sweetclover. Shrubs were
absent.
Urban (1.5%) was restricted to the townsite of Colstrip. Colstrip was
designed such that natural areas and open spaces intermixed with commercial
and residential areas (Aasheim 1980). As a result, roughly 25% of the townsite
consisted of native pine savannah and riparian vegetation-cover types. In
developed portions, grasses, including brome grasses and Kentucky bluegrass,
and trees and shrubs, including ponderosa pine, Rocky Mountain juniper, and
crabapple (Pvrus spp.). were common. Forb diversity and abundance was low in
developed areas, and occurrence was limited primarily to non-paved disturbed
sites and urban-pine savannah ecotones.
Spatial Distribution of Deer
Deer exhibited a clumped distribution during 8 fixed-wing aerial survey
flights, occurring in only 44.8% of the 910 grid cells comprising the study area
(Figure 5). Deer distribution was least clumped during summer (21.8% cell
occupation) and most clumped during winter (10.3% cell occupation) (Figures 6
and 7). Fall and spring dispersion patterns were intermediate with 17.1 and
13.1% of available cells occupied, respectively (Figures 8 and 9). In general,
deer were most abundant in areas where contiguous blocks of pine savannah
extended along slopes and tops of major ridges adjacent to active mining and
reclamation. These areas were topographically diverse and provided the rough
36
terrain favored by mule deer (Severson 1981) as well as abundant hiding and
thermal cover throughout the year. They also provided succulent forage during
summer.
Kilometers
+V *
+
\
+
Ridgetop
i R eclam ation
!8888883 Colstrip
+ D e e r group
'
Figure 5. Distribution of mule deer groups on the Rosebud Mine / Colstrip aerial
survey area based on 8 fixed-wing aerial surveys (n = 2 1 season), summer 1992 spring 1994.
Deer concentrations were lowest in areas distant to mining disturbance
and reclamation, regardless of habitat characteristics. These areas, for example
the northeast and southwest portions of the survey area, because they were
removed from active mining sites, generally received relatively intense grazing
pressure by domestic livestock. Livestock may impact deer negatively by
removing preferred forage species (Mackie 1970, 1976, and 1978, Kie et al.
1991, Loft et al. 1991), reducing hiding cover for fawns (Bowyer and Bleich 1984,
Bowyer 1986, Loft et al. 1987), shifting deer distributions to more energetically
37
W
--------E
+
++
Rldgetop
I
I Reclamation
38888881 Colstrlp
+ Deer group
Figure 6. Summer distribution of mule deer groups on the Rosebud Mine /
Colstrip aerial survey area based on 2 fixed-wing aerial surveys conducted during
July 1992 and 1993.
'
Ridgetop
I
I Reclamation
KSSSSfii Colstrip
+ Deer group
Figure 7. Winter distribution of mule deer groups on the Rosebud Mine / Colstrip
aerial survey area based on 2 fixed-wing aerial surveys conducted during
December 1992 and February 1993.
38
+
+ + + k t.
Rldgetop
I
I R eclam ation
KB&ABB Colstrlp
+ D e e r group
Figure 8. Fall distribution of mule deer groups on the Rosebud Mine / Colstrip
aerial survey area based on 2 fixed-wing aerial surveys conducted during
October 1992 and 1993.
Rldgetop
Reclam ation
BSiSSi Colstrip
+ D e e r group
Figure 9. Spring distribution of mule deer groups on the Rosebud Mine / Colstrip
aerial survey area based on 2 fixed-wing aerial surveys conducted during March
1993 and May 1994.
39
costly habitats (Loft 1988), and altering activity schedules resulting in higher risks
of predation (Kie et al. 1991).
Deer groups (all age / sex classes combined) observed during fixed-wing
aerial surveys occupied grid cells containing specific vegetation-cover types
disproportionate to their availability on the survey area during summer
(X2=18.129, DF=7, P=O.013), fall (X2=27.817, DF=7, P=0.000), and winter
(X2=17.602, DF=3, P=0.016), but not during spring (X2= I 3.962, DF=7, P=0.058)
(Table 4). Grid cells containing reclamation and disturbance types were occupied
more than expected during summer and fall (P<0.05); those containing pine
savannah / urban and agriculture types were occupied less than expected during
'
fall and winter (P<0.05).
Grid cells containing specific vegetation-cover types were used differently
by adult male, productive female (females with fawns), and non-productive female
(females without fawns) groups. Adult male groups occupied grid cells containing
specific vegetation-cover types disproportionate to availability (during summer
(X2=26.748, DF=7, P=0.001), fall (X2=52.272, DF=7, P=0.000), and winter
(X2=20.386, DF=7, P=0.005). Specific selection, however, was exhibited only
during summer when grid cells containing urban types were occupied less than
expected (P<0.05) (Table 5).
Productive and non-productive female groups also occupied grid cells
containing specific vegetation-cover types disproportionate to availability during
summer (non-productive: X2=34.835, DF=7, P=0.000; productive: X2=27.978,
DF=7, P=O-OOO), fall (non-productive: X 2=28.516, DF=7, P=0.000; productive:
40
Table 4. Proportional availability and use of grid cells containing specific
vegetation-cover types by mule deer (all age / sex classes), Rosebud Mine /
Colstrip, Montana, summer 1992-spring 1994.
Season3
SUM
GL9
MX
PS
RP
DS
RC
AG
UR
FAL
GL
MX
PS
RP
DS
RC
AG
UR
W NT
GL
MX
PS
RP
DS
RC
AG
UR
SPR
GL
MX
PS
RP
DS
RC
AG
UR
Ob
Ec
Nd
X2
Confidence interval6
Selectionf
0.283
0.717
0.665
0.613
0.377
0.325
0.120
0.042
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
191
191
191
191
191
191
191
191
2.620
0.540
0.000
0.200
7.389
5.889
0.658
0.833
0.194
0.628
0.571
0.516
0.281
0.232
0.056
0.002
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.372
0.806
0.759
0.710
0.473
0.418
0.184
0.082
NS
NS
NS
NS
P
P
NS
NS
0.297
0.690
0.516
0.581
0.381
0.323
0.155
0.006
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
155
155
155
155
155
155
155
155
3.705
1.082
3.378
0.712
7.837
5.681
3.789
1.633
0.196
0.588
0.406
0.472
0.274
0.220
0.075
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.398
0T92
0.626
0.690
0.488
0.426
0.235
0.023
NS
NS
A
NS
P
P
NS
A
0.191
0.660
0.564
0.574
0.319
0.330
0.032
0.021
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
94
94
94
94
94
94
94
94
0.155
1.903
1.562
0.867
2.377
6.427
4.178
0.133
0.080
0.526
0.424
0.434
0.187
0.197
0.000
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.302
0.794
0.704
0.714
0.451
0.463
0.082
0.062
NS
NS
NS
NS
NS
NS
A
NS
0.294
0.672
0.605
0.613
0.319
0.269
0.118
0.059
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
119
119
119
119
119
119
119
119
3.457
1.547
0.559
, 0.200
2.377
1.472
0.557
3.793
0.179
0.554
0.482
0.491
0.202
0.157
0.037
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.409
0.790
0.728
0.735
0.436
0.381
0.199
0.118
NS
NS
NS
NS
NS
NS
NS
NS
41
Table 4. Continued, 2 of 2.
a Seasonal designations: SUM=summer, 01 June-31 August; FAL=fall, 01
September-30 November; WNT=winter, 01 December-28 February; and
SPR=spring, 01 March-31 May. Use of grid cells containing specific
vegetation-cover types was disproportionate to availability during summer
(X2= 18.129, DF=7, £=0.013), fall (X2=27.817, DF=7, £=0.000), and winter
(X2= I 7.602, DF=3, £=0.016), but not spring (X2= I 3.962, DF=7, £=0.058).
b The proportion of deer groups observed during aerial surveys occupying grid
cells containing the vegetation-cover type of interest. This proportion
represents observed use of grid cells.
c The proportion of grid cells comprising the study area containing the
vegetation-cover type of interest. This proportion represents expected use of
grid cells.
d The number of grid cells in which deer were observed during survey flights.
e 95% Bonferroni confidence interval constructed about the observed proportion
of grid cell use.
f Selection of vegetation-cover types: NS denotes no selection; P denotes
preference; and A denotes avoidance. Preference is indicated when the
expected proportion of grid cell use is less than the lower confidence limit of
observed use (£<0.05), avoidance is indicated when the expected proportion of
grid cell use is greater than the upper confidence limit of observed use
(£<0.05), and no selection is indicated when the expected proportion of grid cell
use falls within the confidence limits of observed use (£ > 0 .0 5 ).'
9 Designation of vegetation-cover types: GL=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
42
Table 5. Proportional availability and use of grid cells containing specific
vegetation-cover types by adult male mule deer, Rosebud Mine / Colstrip1
Montana, summer 1992-winter 1993.
Season3
SUM
GL9
MX
PS
. RP
DS
RC
AG
UR
FAL
GL
MX
PS
RP
DS
RC
AG
UR
W NT
GL
MX
PS
RP
DS
RC
AG
UR
Ob
Ec
Nd
X2
Confidence interval® Selectionf
0.200
0.677
0.677
0.492
0.369
0.369
0.123
0.000
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
65
65
65
65
65
65
65
65
0.039
1.410
0.018
3.798
6.533
11.425
0.825
2.700
0.064
0.518
0.518
0.322
0.205
0.205
0.011
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.336
0.836
0.836
0.662
0.533
0.533
0.235
0.000
NS
NS
NS
NS
NS
NS
NS
A
0.368
0.667
0.509
0.561
0.386
0.246
0.140
0.105
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
57
57
57
57
57
57
57
57
12.096
1.691
3.701
1.193
8.415
0.511
2.132
22.533
0.193
0.496
0.327
0.381
0.209
0.090
0,014
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.543
0,838
0.691
0.741
0.563
0.402
0.266
0.216
NS
NS
NS
NS
NS
NS
NS
NS
0.194
0.710
0.516
0.677
0.290
0.323
0.032
0.065
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
31
31
31
31
31
31
31
31
0.108
0.663
3.378
0.121
0.909
5.681
4.178
5.348
0.000
0.486
0.270
0.447
0.066
0.093
0.000
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.389
0.934
0.726
0.907
0.514
0.553
0.119
0.186
NS
NS
NS
NS
NS
NS
NS
NS
a Seasonal designations: SUM=summef, 01 June-31 August; FAL=fall, 01
September-30 November; and WNT=winter, 01 December-28 February. Use
of grid cells containing specific vegetation-cover types was disproportionate to
availability during summer (X2=26.748, DF=7, P=0.001), fall (X2=52.272, DF=7,
P=0.000), and winter (X2=20.386, DF=7, P =0.005).
b The proportion of adult male groups observed during aerial surveys occupying
grid cells containing the vegetation-cover type of interest. This proportion
represents observed use of grid cells.
c The proportion of grid cells comprising the study area containing the
vegetation-cover type of interest. This proportion represents expected use of
grid cells.
<
43
Table 5. Continued, 2 of 2.
d The number of grid cells in which adult males were observed during survey
flights.
e 95% Bonferroni confidence interval constructed about the observed proportion
of grid cell use.
f Selection of vegetation-cover types: NS denotes no selection; P denotes
preference; and A denotes avoidance. Preference is indicated when the
expected proportion of grid cell use is less than the lower confidence limit of
observed use (P<0.05), avoidance is indicated when the expected proportion of
grid cell use is greater than the upper confidence limit of observed use
(P^O.05), and no selection is indicated when the expected proportion of grid cell
use falls within the confidence limits of observed use (P>0.05).
9 Designation of vegetation-cover types: GL=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
44
X2=40.949, DF=7, P=0.000), and winter (non-productive: X2=28.343, DF=7,
P=O-OOO; productive: X 2=68.905, DF=7, P=0.000). Groups containing non­
productive females used grid cells containing disturbance and reclamation types
more than expected during summer and those containing urban types less than
expected during fall and winter (P<0.05) (Table 6). Contrastingly, groups
containing productive females exhibited no type-specific selection during summer,
used grid cells containing pine savannah and disturbance / reclamation types less
than and greater than expected, respectively during fall, and Used grid cells
containing reclamation types more than expected during winter (P<0.05) (Table
7,
'
Although differential use of space and vegetation by males and females is
widely documented among mule deer (Dasmann and Taber 1956, Miller 1970,
Boukhout 1972, King and Smith 1980, Bowyer 1984, Ordway and Krausman
1986, Scarbrough and Krausman 1988, Hamlin and Mackie 1989, Wood et al.
1989, Main and Coblentz 1991, Pac et al. 1991, Main 1994a), differences
between reproductive classes of females has been less studied. Use of
vegetation-cover types by productive and non-productive female groups differed
mainly in early summer when relatively open reclamation types were used by non­
productive female groups but avoided by productive female groups, possibly
because fawn concealment cover often was lacking. As summer progressed,
however, fawns likely became less dependent on a hiding strategy as well as
more mobile (de Vos et al. 1967), allowing productive females to increase use of
reclamation.
45
Table 6. Proportional availability and use of grid cells containing specific
vegetation-cover types by adult female mule deer without fawns, Rosebud Mine /
Colstrip, Montana, summer 1992-winter 1993.
Season3
SUM
GL9
MX
PS
RP
DS
RC
AG
UR
FAL
GL
MX
PS
RP
DS
RC
AG
UR
WNT
GL
MX
PS
RP
DS
RC
AG
UR
Ob
Ec
Nd
X2
Confidence interval6
Selectionf
0.312
0.706
0.642
0.633
0.431
0.367
0.147
0.037
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
109
109
109
109
109
109
109
109
5.076
0.739
0.086
0.039
14.545
11.134
2.846
0.370
0.190
0.586
0.516
0.506
0.301
0.240
0.054
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.434
0.826
0.768
0.760
0.561
0.494
0.240
0.087
NS
NS
NS
NS
P
P
NS
NS
0.308
0.808
0.577
0.615
0.385
0.269
0.192
0.000
0.209
0.782
0.666
0.469
0.243
0.213
0.095
0.027
26
26
26
26
26
26
26
26
4.689
0.086
1.189
0.178
8.298
1.472
9.904
2.700
0.060
0.596
0.311
0.353
0.123
0.030
0.000
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.556
1.000
0.843
0.877
0.647
0.508
0.404
0.000
NS
NS
NS
NS
NS
NS
NS
A
0.291
0.727
0.564
0.673
0.418
0.327
0.055
0.000
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
55
55
55
55
55
55
55
55
3.217
0.387
1.562
0.089
12.603
6.101
1.684
2.700
0.123 <
0.562 <
0.381 <
0.499 <
0.236 <
0 .1 5 3 <
0.000 <
0.000 <
O
O
O
O
O
O
O
O
< 0.459
< 0.892
< 0.747
< 0.847
< 0.600
< 0.5 01
< 0.139
< 0.000
NS
NS
NS
NS
NS
NS
NS
A
a Seasonal designations: SUM=summer, 01 June-31 August; FAL=fall, 01
September-30 November; and WNT=winter, 01 December-28 February. Use
of grid cells containing specific vegetation-cover types was disproportionate to
availability during summer (X2=34.835, DF=7, P=O OOO), fall (X2=28.516, DF=7,
E=0.000), and winter (X2=28.343, DF=7, P=0.000).
b The proportion of female groups without fawns observed during aerial surveys
occupying grid cells containing the vegetation-cover type of interest. This
proportion represents observed use of grid cells.
c The proportion of cells comprising the study area containing the vegetationcover type of interest. This proportion represents expected use of cells.
,
.
46
Table 6. Continued, 2 of 2.
d The number of grid cells in which females without fawns were observed during
survey flights.
e 95% Bonferroni confidence interval constructed about the observed proportion
of grid cell use.
f Selection of vegetation-cover types: NS denotes no selection; P denotes
preference; and A denotes.avoidance. Preference is indicated when the
expected proportion of grid cell use is less than the lower confidence limit of
observed use (P<0.05), avoidance is indicated when the expected proportion of
grid cell use is greater than the upper confidence limit of observed use
(P<0.05), and no selection is indicated when the expected proportion of grid cell
use falls within the confidence limits of observed use (P>0.05).
9 Designation of vegetation-cover types: GL=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
47
Table 7. Proportional availability and use of. grid cells containing specific
vegetation-cover types by adult female mule deer with fawns, Rosebud Mine /
Colstrip1 Montana, summer 1992-rwinter 1993.
Season3
SUM
GL9
MX
PS
RP
DS
RC
AG
UR
FAL
GL
MX
PS
RP
DS
RC
AG
UR
WNT
GL
MX
PS
RP
DS
RC
AG
UR
Ob
Ec
Nd
X2
Confidence interval6
Selectionf
0.310
0.762
0.643
0.643
0.333
0.286
0.095
0.095
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
42
42
42
42
42
42
42
42
4.881
0.051
0.079
' 0.006
3.333
2.502
0.000
17.126
0.114
0.582
0.440
0.440
0.134
0.095
0.000
0.000
<G<
< O <
< O <
< O <
< O <
< O <
< O <
< O <
0.506
0.942
0.846
0.846
0.532
0.477
0.219
0.219
NS
NS
NS
NS
NS
NS
NS
NS
0.301
0.660
0.524
0.583
0.398
0.359
0.136
0.078
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
103
103
103
103
103
103
103
103
4.050
1.903
3.028
0.671
9.887
10.008
1.769
9.633
0.177
0.532
0.389
0.450
0.266
0.229
0.043
0.006
<
<
<
<
<
<
<
<
0
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.425
0.788
0.659
0.716
0.530
0.489
0.229
0.150
NS
NS
A
NS
P
P
NS
NS
0.097
0.645
0.645
0.516
0.290
0.484
0.032
0.097
0.209
0.782
0.666
0.649
0.243
0.213
0.095
0.027
31
31
31
31
31
31
31
31
6.002
2.400
0.066
2.726
0.909
34.479
4.178
18.148
0.000
0.409
0.409
0.270
0.066
0.238
0.000
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.243
0.881
0.881
0.762
0.514
0.730
0.119
0.243
NS
NS
NS
NS
NS
P
NS
NS
a Seasonal designations: SUM=Summer, 01 June-31 August; FAL=fall, 01
September- 30 November; and WNT=winter, 01 December-28 February. Use
of grid cells containing specific vegetation-cover types was disproportionate to
availability during summer (X2=27.978, DF=7, £=0.000), fall (X2=40.949, DF=7,
P=0.000), and winter (X2=68.905, DF=7, £=0.000).
b The proportion of female groups with fawns observed during aerial surveys
occupying grid cells containing the vegetation-cover type of interest. This
proportion represents observed use of grid cells.
c The proportion of grid cells comprising the study area containing the
vegetation-cover type of interest. This proportion represents expected use of
grid cells.
48
Table 7. Continued, 2 of 2.
d The number of grid cells in which females with fawns were observed during
survey flights.
e 95% Bonferroni confidence interval constructed about the observed proportion
of igrid cell use.
f Selection of vegetation-cover types: NS denotes no selection; P denotes
preference; and A denotes avoidance. Preference is indicated when the
expected proportion of grid cell use is less than the lower confidence limit of
observed use (P<0.05), avoidance is indicated when the expected proportion of
grid cell use is greater than the upper confidence limit of observed use
(P<0.05), and no selection is indicated when the expected proportion of grid cell
use falls within the confidence limits of observed use (P>0,.05).
9 Designation of vegetation-cover types: GL=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
49
Male and female deer groups were distributed differently with respect to
the number of vegetation-cover types contained within occupied grid cells. Grid
cells selected by female groups contained more vegetation-cover types than cells
not used during summer (X2=28.218, D F = I1 P=0.001) and fall (X2=4.998, DF=1,
P=0.025), but not duripg winter (X2= 1.269, D F=1, P=0.260). No such relationship
was indicated for male groups during summer (X2=O.949, DF=1, P=0.758), fall
(X2= 1.108, DF=1, P=0.293), or winter (X2=0.691, DF=1, P=0.406). Number of
vegetation-cover types within grid cells occupied by female groups exceeded that
of cells occupied by male groups during summer (X2=6.590, DF=1, P=0.010), but
not during fall (X2=O. 115, DF=1, P=0.734) or winter (X2=1.377, DF=1, P=0.241).
Finally, the number of vegetation-cover types within grid cells occupied by female
groups differed seasonally (X2=7.016, DF=2, P=0.030), mean number of types /
I
cell during summer exceeding that during winter (P < 0.05). The number of types
within cells occupied by male groups did not differ seasonally (X2=1.640, DF=2,
P=0.440), Thus, male and female group distributions differed most with respect
to diversity of vegetation-cover types within occupied grid cells during summer
and fall and least during winter.
Selection for grid cells containing different vegetation-cover types resulted
in distribution differences between male and productive female groups and
productive and non-productive female groups, but not between male and non­
productive female groups. During summer, adult male and non-productive female
groups were distributed randomly with respect to each other among grid cells
(X2=O.604, DF= 1, P=0.457), sharing 29 of 67 possible cells. However, of 67 grid
50
cells potentially occupied by both adult male and productive female groups, only 9
actually were shared, indicating non-random, segregated distributions
(X2=17.918, DF=1, P<0.001). Further, only 10 of 46 grid cells potentially
occupied by both productive and non-productive female groups were shared,
again indicating segregation (X2=7.348, DF=1, P=0.008). During fall, adult male
groups were distributed randomly with respect to both non-productive (X2=O.009,
DF=1, P=0.920) and productive (X2=0.711, DF=1, P=0.426) female groups.
However, productive and non-productive female groups continued to maintain
disparate distributions, as only 8 of 36 possible cells were shared (X2=5.556,
DF=1, P=0.015). During winter, adult male and non-productive female groups
exhibited a non-random, clumped distribution (X2=4.661, DF=1, P =0.035)
occurring together in 24 of 31 possible grid cells. Productive female groups,
however, reestablished a separate distribution from adult male groups, as only 3
of 31 possible grid cells were shared (X2=10.081, DF=T, P =0.004). Productive
female groups also maintained segregation from non-productive female groups,
sharing only 3 of 31 possible grid cells (X2= 10.081, DF=1, P =0.001).
Male and non-productive female deer tend to occur in common areas
separate from those occupied by productive females (Glutton-Brock et al. 1982,
Hamlin and Mackie 1989).. For females to successfully rear offspring, they must
occupy diverse habitats that provide succulent forage throughout the lactation
period, isolation from other deer and / or ungulate species, and security from
predators (Dusek et al. 1989, Wood et al. 1989, Pac et al. 1991). Females
occupying such high-quality, diverse, “reproductive” habitats tend to successfully
51
rear fawns and maintain residency yearlong, resulting in territorial expansion of
the matrilineal group as reproductive habitats surrounding that of the matriarch
are occupied and maintained by female offspring ((Dzoga et al. 1982, Pac et al.
1991, Porter et al. 1992). As available reproductive habitats are filled, however,
females entering the population, either through birth or immigration, must occupy
more marginal habitats, generally peripheral to reproductive habitats, where the
probabilities of successful reproduction and yearlong residency are reduced
(Hamlin and Mackie 1989). Thus, productive and non-productive females exhibit
different distributions not because of disparate survival / reproductive strategies,
but because habitats occupied by each determine reproductive success or failure.
In contrast, males are inherently more mobile than females, ranging over
larger areas to survive and reproduce (Marchinton and Hirth 1984, Hamlin and
Mackie 1989, Main 1994a). Because of greater mobility and the physiological
capacity to exist in habitats providing lesser forage quality (Glutton-Brock et al.
1982, Beier 1987, Putman 1988, Main 1994a), males tend to defer reproductive
habitats to females, which, upon establishing home ranges within these habitats
via either immigration or expansion of matrilineal groups, maintain a constant
presence. Thus, males are left to occupy less diverse, more marginal
maintenance habitats that support survival of adults but not production of fawns.
These habitats, peripheral to reproductive habitats, also are occupied by non­
productive females; thus, distributions of males and non-productive females may
be similar.
The overall distribution of deer during this study reflected inherent
52
behavioral differences between males and females as well as differences in
female reproductive success imposed by habitat. Males and non-productive
females exhibited similar distributions that differed from the distribution of
reproductive females during most seasons. Females occupied areas with a
greater diversity of vegetation-cover types than did males during the summer
fawning season. This suggested greater fill of diverse reproductive habitats by
females and use of more marginal habitats by males. In summer, non-productive
females tended to use areas with more open vegetation and less topographic
diversity than selected by productive females, suggesting perhaps that these
habitats lacked elements necessary for high fawn survival.
General Movement Patterns
Two general movement patterns, yearlong residency and seasonal
migration, were identified among 55 radiocolIared yearling and adult mule deer.
Most deer, 39 of 44 (88.6% ) adult females and all adult males (n=11, 100.0%),
were yearlong residents. Although these deer often shifted areas of intensive
use seasonally, shifts were not so extreme that distinct seasonal home ranges
could be identified.
Seasonal migrants exhibited spring and fall / winter movements between
distinct summer and winter home ranges and were relatively uncommon
compared to other deer populations in Montana (lhsle 1982, Wood 1986, Hamlin
and Mackie 1989, Jackson 1990, Pac et al. 1991, Stansberry 1991). Only 5 of
the 55 radiocollared adults (9.1%), all females, exhibited seasonal migration.
53
Migration to summer ranges occurred during late April / early May, whereas
timing of return migration to winter ranges was more variable, occurring
September-December. Limited evidence suggested that timing of winter
migration may have been influenced by the fate of fawns born on summer ranges.
O f 7 winter migration events for which the productivity of females could be
assessed, the mean dates of return to winter ranges were 15 September for 2
non-productive females and 13 December for 5 reproductive females. Migration
movements were directionally uniform, with all 5 migrants moving west-southwest
to summer home ranges in the vicinity of the Sarpy Creek Coal Mine. Mean
migration distance was 24.2±4.0 (SE) km.
Migrants wintered in 2 discrete areas
on the Rosebud Mine separated by approximately 14.6 km.
^
Accessory areas, sites located outside the boundaries of normal seasonal
home ranges (Pac et al. 1991), were used seasonally by 4 resident females.
Three occupied accessory areas only during extended periods of winter cold
coupled with above average snow fall. The other occupied an accessory area
prior to and during early fawn-rearing in June and July. Use of accessory areas
likely was necessitated by the need for females to either (1) satisfy temporary
resource deficiencies within normal seasonal home ranges (e.g., forage or
thermal cover deficiencies encountered during periods of weather extremes), or
(2) avoid aggression from dominant females during the early fawn-rearing period
(Pac et al. 1991). Accessory areas were used by females for relatively short
periods of time (m ean=33.8±18.3 (SE) days), after which they returned to normal
seasonal home ranges, presumably as soon as climatic, forage, and / or social
54
conditions allowed.
Movement patterns of deer generally reflect the ability of the environment
to provide for individual needs (Wood et al. 1989, Pac et al. 1991). Deer
inhabiting diverse, stable, high-quality habitats tend to exhibit yearlong residency
patterns and have reduced need of accessory areas. Alternatively, deer
occupying less diverse habitats in which resources are widely distributed or more
variable through space or time exhibit greater movement or migration between
distinct seasonal home ranges capable of temporarily supporting deer survival
and / or reproduction. These deer also are more likely to use accessory areas
either during migration or while occupying seasonal home ranges. The high
proportion of yearlong resident deer and infrequent use of accessory areas by
deer on the Rosebud Mine / Colstrip study area suggested (1) high overall quality
and diversity of habitats capable of satisfying yearlong resource requirements of
deer, and / or (2) deer population density below that where individual use of lowdiversity, marginal habitats and seasonal movements to and from more suitable
areas is required.
Home Range. Mobility, and Activity
Home range characteristics differed between females and males and
among seasons. Annual home range size for males averaged 1,533.7+513.2
(SE) ha, more than double the 545.7±320.8 (SE) ha mean for females (rank
sum=140.0, P=O.003) (Table 8). Home range size differed seasonally for females
(X2= 10.234, DF=3, P=0.017), spring home range sizes exceeding those in
55
summer (P<0.05), but not for males (X2=O.325, DF=3, P=0.955) (Table 8).
Seasonal home ranges of males were larger than those of females during
summer (rank sum=803.0, P=0.001), fall (rank sum=324.0, P =0.007), and spring
(rank sum=119.0, P=0.048), but not during winter (rank sum =110.0, P=0.314).
Diel home range size, obtained for females only, averaged 42.7+24.3 (SE) ha
across all seasons. Sizes differed seasonally (X2=32.730, DF=3, P=0.000),
spring (84.3±110.5 (SE) ha) and fall (52.4+27.6 (SE) ha) home range sizes
exceeding those during summer (27.7±25.2 (SE) ha) and winter (21.6±15.4 (SE)
ha) (P<0.05) (Table 8).
Table 8. Mean annual, seasonal, and diel home range sizes (ha) for
radiocollared adult mule deer, Rosebud Mine / Colstrip, Montana, fall 1992-spring
1994.
Period
Season
Annual3
Mean home
range size+SE ( c f )
Mean home
range size+SE (?)
1533.7±513.2 (4)b
545:7±320.8 (34)
348.8±176.4
408.1±196.0
323.2±219.5
399.9±232.2
126.7±103.5
155.6±140.8
192.4±154.2
205.6±158.5
Seasonal0
Summer
Fall
Winter
Spring
(12)
(07)
(04)
(04)
(69)
(47)
(38)
(34)
Diel
Summer
Fall
Winter
Spring
NAd
NA
NA
NA
27.7±25.2
52.4±27.6
21.6±15.4
84.3±98.5
(51 )e
(24)
(26)
(26)
a 01 June 1993-31 May 1994.
b The number of deer from which estimates were derived.
c Seasonal designations: summer, 01 June-30 August; fall, 01 September-30
November; winter, 01 December-28 February; and spring, 01 March-31 May.
d Estimates not available for adult male deer.
e The number of diel monitoring sessions from which estimates were derived.
56
Despite both the similarity of movement patterns exhibited by most deer on
the study area (i.e., yearlong residency of home ranges with little use of
accessory areas) and the general sex-specific, seasonal, and diel trends
identified, home range size and configuration varied substantially among
individuals. Presumably, this resulted from unique sociological and physiological
characteristics of each deer and the fixed and variable components of the
particular areas or habitat it occupied (Hamlin and Mackie 1989, Pac et al. 1991).
Deer mobility, determined by total distance traveled over a 24-hour period,
differed seasonally for adult females (X2=24.172, DF=3, P=O. 000) and paralleled
trends for seasonal home range size. Distances traveled during summer, fall,
and spring averaged 6.6+3.2, 7.7±2.0, and 7.4±3.1 (SE) km, respectively, and
each exceeded the 4.7+1.7 (SE) km mean for winter (P<0.05). Mobility,
determined by meters traveled between successive radiolocations, differed
among the 4 diel periods during summer (X2= 18.742, DF=3, P=0.000), winter
(X2= 14.272, DF=3, P=0.003), and spring (X2= 11.993, DF=3, P=0.007), but not
during fall (X2=5.247, DF=3, P =0.155) (Table 9). Females were most mobile
during nocturnal, afternoon / evening, and diurnal hours during summer, winter,
and spring, respectively.
Diel activity rates differed seasonally (X2= 13.025, DF=3, P=0.005) and
matched trends established for mobility. During summer, fall, and spring, females
were active on average during 46.8±21.7, 48.0±13.4, and 46.3+19.5 (SE) %,
respectively, of 24-hour periods. Each of these means exceeded the 38.1 ±14.2
(SE) % mean for winter (P<0.05). In contrast to mobility, activity differed among
57
Table 9. Mean (±SE) number of meters traveled per hour by radiocollared adult
female mule deer during die! periods 1-4, Rosebud Mine / Colstrip, Montana,
summer 1992-spring 1994. '
Season3
Summer
Fall
Winter
Spring
Nb
Period-13
Period-2
Period-3
Period-4
X2
265.6(AB)d
±113.4
298.1 (A)
470
±143.2
123.8 (B)
519
±107.3
542
348.1 (A)
±221.9
218.4(B)
±144.3
379.3(A)
±212.4
215.0(A)
±184.3
316.8(A)
±141.7
308.7(A)
±189.8
291.4(A)
±100.1
252.3(A)
±133.5
218.3(B)
±113.4
294.9(A)
±104.0
261.3(A)
±221.6
161.9(B)
±104.2
215.7(B)
±106.8
18.742*
1022
5.247
14.272*
11.993*
* Denotes statistical significance (P<0.05).
a Seasonal designations: summer, 01 June-31 August; fall, 01 September-30
November; winter, 01 December-28 February; spring, 01 March-31 May.
b The number of mobility estimates obtained from diel monitoring used to derive
mean values for each diel period. Individual estimates were obtained by
measuring the minimum straight-line distance (m) traveled between successive
locations obtained at I -hour intervals during each diel period of a monitoring
session.
'
0 Diel period designations: period-1, sunrise to mid day; period-2, mid day to
sunset; period-3, sunset to mid night; and period-4, mid night to sunrise.
d Mean mobility estimates (meters traveled per hour). Capital letters in
parentheses represent the results of multiple comparison tests. Mean values
with the same letters did not differ (P>0.05). Mean values with different letters
differed (P<0.05).
,
’
diel periods only during winter (X2=22.219, DF=3, P=0.000), when activity during
diel periods 2 (mid day to sunset) and 3 (sunset to mid night) exceeded that
during period 4 (mid night to sunrise) (P<0.05) (Table 10).
Patterns of mobility and activity, also varied substantially among individual
deer. Like home range size and shape, the patterns apparently reflected
individual strategies to cope with local habitat conditions. Despite the variability,
seasonal and diel trends identified also likely reflected strategies to maximize
58
Table 10. Mean (±SE) percentage of radiocollared adult female mule deer
activity during die! periods 1-4, Rosebud Mine / Colstrip, Montana, summer 1992spring 1994.
Season3
Summer
Fall
Nb
136
96
Winter
136
Spring
84
Period-13
Period-2
Period-3
Period-4
X2
44.7 (A)d
±27.3
43.3 (A)
±19.4
35.0(AB)
±29.8
46.9 (A)
±33.8
43.0(A)
±24.1
56.1(A)
±27.9
48.9(A)
±22.7
52.1(A)
±42.6
50.9(A)
±29.3
47.0(A)
±17.3
45.8(A)
. ±18.6
42.1(A)
±29.6
48.6(A)
±31.7
45.7(A)
±21.3
22.7(B)
±21.5
44.1(A)
±45.7
2.877
4.345
22.219*
4.463
* Denotes statistical significance (P<0.05).
a Seasonal designations: summer, 01 June-31 August; fall, 01 September-30
November; winter, 01 December-28 February; spring, 01 March-31 May.
b The number of diet monitoring sessions from which period-specific activity
estimates were derived.
c Diel period designations: period-1, sunrise to mid day; period-2, mid day to
sunset; period-3, sunset to mid night; and period-4, mid night to sunrise.
d Mean activity estimates (the percentage of active locations obtained in a given
diel period during a die! monitoring session). Capital letters in parentheses
represent the results of multiple comparison tests. Mean values with the same
letters did not differ (P>0.05). Mean values with different letters differed
(P<0.05).
female productivity during summer and survival during winter. Mobility and
activity were much reduced during winter, supporting the concept that deer
enhance survival probabilities during winter by reducing metabolism (Mautz et al.
1985, Parker 1994) when forage quality is low (Cook 1972) and time / energy
required to digest forage is high (Arnold 1985, Welch and Hooper 1988, Parker
1994).
Mobility and activity patterns of females over diel cycles also varied
seasonally such that energy loss associated with movement was minimized and
59
reproductive success and probability of survival were maximized. For example,
during summer, females were more mobile and active nocturnally than diurnally,
thereby reducing energetic costs associated with dissipating heat generated by
movements during high-temperature periods (Short 1981, Parker 1994).
Alternatively, during winter and spring, females were more mobile and active
diurnally than nocturnally, thereby reducing energetic costs associated with heat
loss while active during cold nocturnal periods ((Dzoga and Verme 1970, Short
1981, Mautz e ta l. 1985).
Use of Vegetation-Cover Types bv Radiocollared Deer
Use of vegetation-cover types was disproportionate to availability for
radiocollared adult male and female deer during each season at both levels of
analysis (Tables 11-14). At the second-order (study area) level, mixed shrub,
agriculture, and urban types generally were included in home ranges of both
males and females less than was expected. At the third-order (home range)
level, females always used pine savannah and riparian types more than
expected, often used grassland, reclamation, and urban types more than
expected, and always used mixed shrub and disturbance types less than
expected. Males generally used pine savannah types more than expected and
mixed shrub, disturbance, and urban types less than expected.
Seasonal use of vegetation-cover types differed subtly between males and
females. Female use of types providing concealment cover was high during
summer, as would be predicted of females with young fawns (King and Smith
60
Table 11. Proportional occurrence of vegetation-cover types within home ranges
of radiocollared adult female mule deer in relation to proportional availability of
vegetation-cover types within the study area, Rosebud Mine / Colstrip, Montana,
summer 1993-spring 1994.
Season3
Nb
Oc
Ed
Confidence interval6
Selection'
SUM
GL9
MX
PS
RP
DS
RC
AG
UR
36
36
36
36
36
36
36
36
0.083
0.249
0.183
0.041
0.202
0.209
0.019
0.013
0.074
0.311
0.167
0.035
0.188
0.181
0.028
0.015
0.072 <
0.232 <
0.168 <
0.033 <
0.186 <
0 .1 9 3 <
0.014 <
0.009 <
<
<
<
<
<
<
<
<
0.094
0.266
0.198
0.049
0.218
0.225
0.024
0.017
NS
A
P
NS
NS
P
A
NS
GL
MX
PS
RP
DS
RC
AG
UR
36
36
36
36
36
36
36
36
0.078
0.238
0.113
0.034
0.194
0.316
0.024
0.004
0.074
0.311
0.167
0.035
0.188
0.181
0.028
0.015.
0.069 < O <
0.223 < O <
0.102 < O <
0.028 < O <
0 .1 8 0 < O <
0.300 < O <
0.019 < O <
0.002 < O <
0.087
0.253
0.124
0.040
0.208
0.332
0.029
0.006
NS
A
A
NS
NS
P
NS
A
GL
MX
PS
RP
DS
RC
AG
UR
38
38
38
38
38
38
38
38
0.083
0.251
0.164
0.038
0.190
0.252
0.015
0.008
0.074
0.311
0.167
0.035
0.188
0.181
0.028
0.015
0.074 < O < 0.092
0.237 < O < 0.265
0.152 < O < 0.176
0.032 < O < 0.044
0 .1 7 7 < O < 0.203
0.238 < O < 0.266
0.011 < O < 0.019
0 .0 0 5 < O < 0.0 11
NS
A
NS
NS
P
P
A
A
GL
MX
PS
RP
DS
RC
AG
UR
34
34
34
34
34
34
34
34
0.136
0.238
0.156
0.033
0.218
0.186
0.028
0.005
0.074
0.311
0.167
0.035
0.188
0.181
0.028
0.015
0.125 <
0.224 <
0.144 <
0.027 <
0.204 <
0 .1 7 3 <
0.023 <
0.003 <
P
A
NS
NS
P
NS
NS
A
O
O
O
O
O
O
O
O
FAL
WNT
SPR
'
O <
O <
O <
O <
O <
0<
O <
O <
0.147
0.252
0.168
0.039
0.232
0.199
0.033
0:007
I
61
Table 11. Continued, 2 of 2.
a Seasonal designations: SUM=summer, 01 June-31 August; FAL=fall, 01
September-30 November; WNT=Winter1 01 December-28 February; and
SPR=spring, 01 March-31 May. Overall use of vegetation-cover types was
disproportionate to availability during summer (X2=121.6, DF=7, £=0.000), fall
(X2=919.5, DF=7, P=O-OOO), winter (X2=359.5, DF=7, P =0.000), and spring
(X2=577.6, DF=7, P=0.000).
b The number of adult females from which selection of vegetation-cover types
was derived.
c The observed proportion of vegetation-cover type use.
d The expected proportion of Vegetation-cover type use.
e 95% Bonferroni confidence interval constructed about the observed proportion
of vegetation-cover type use.
f Expected proportions of vegetation-cover type use less than the lower
confidence limit denote preference (P) (P<0.05), those exceeding the upper
confidence limit denote avoidance (A) (P<0.05), and those falling within the
confidence interval denote lack of selection (NS) (P>0.05).
9 Designation of vegetation-cover types: GL=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
62
Table 12. Proportional occurrence of vegetation-cover types within home ranges
of radiocollared adult male mule deer in relation to proportional availability of
vegetation-cover types within the study area, Rosebud Mine / Colstrip, Montana,
summer 1993-spring 1994.
Nb
Season3
O0
Ed
Confidence interval6
Selection5
SUM
GL9
MX
PS
RP
DS
RC
AG
UR
8
8
8
8
8
8
8
8
0.050
0.381
0.197
0,045
0.120
0.164
0.043
0.000
0.074
0.311
0.167
0.035
0.188
0.181
0.028
0.015
0.039
0.356
0.177
0.034
0.104
0.145
0.033
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.061
0.406
0.217
0.056
0.136
0.183
0.053
0.000
A
P
P
NS
A
NS
P
A
GL
MX
PS
RP
DS
RC
AG
UR
6
6
6
6
6
6
6
6
0.069
0.304
0.090
0.032
0.211
. 0.269
0.022
0.002
0.074
0.311
0.167
0.035
0.188
0.181
0.028
0.015
0.055 <
0.279 <
0.075 <
0.023 <
0 .1 8 9 <
0.245 <
0.014 <
0.000 <
O
O
O
O
O
O
0
O
<
<
<
<
<
<
<
<
0.083
0.329
0.105
0.041
0.233
0.293
0.030
0.004
NS
NS
A
NS
P
P
NS
A
GL
MX
PS
RP
DS
RC
AG
UR
4
4
4
4
4
4
4
4
0.093
0.330
0.098
0.059
0.246
0.127
0.026
0.020
0.074
0.311
0.167
0.035
0.188
0.181
0.028
0.015
0.071
0.294
0.075
0.041
0.213
0.102
0.014
0.009
<G<
< O <
< O <
< O <
< O <
< O <
<O <
< O <
0.115
0.366
0.121
0.072
0.279
0.152
0.038
0.031
NS
NS
A
P
P
A
NS
NS
GL
MX
PS
RP
DS
RC
AG
UR
4
4
4
4
4
4
4
4
0.105
0.244
0.304
0.034
0.088
0.207
0.011
0.007
0.074
0.311
0.167
0.035
0,188
0.181
0.028
0.015
0.084
0.215
0.272
0.022
0.069
0.179
0.004
0.001
<
<
<
<
<
<
<
<
0.126
0.273
0.336
0.046
0.107
0.235
0.018
0.013
P
A
A
NS
A
NS
A
A
FAL
WNT
SPR
.
O <
O <
O <
O <
O <
0<
O <
O <
63
Table 12. Continued, 2 of 2.
a Seasonal designations: SUM=Summer, 01 June-31 August; FAL=fall, 01
September-30 November; WNT=winter, 01 December-28 February; and
SPR=spring, 01 March-31 May. Overall use of vegetation-cover types was
disproportionate to availability during summer (X2=236.5, DF=7, P=0.000), fall
(X2=243.9, DF=7, P=0.000), winter (X2= 112.5, DF=7, P=0.000), and spring
(X2=335.4, DF=7; P=0.000).
b The number of adult males from which selection of vegetation-cover types was
derived.
c The observed proportion of vegetation-cover type use.
d The expected proportion of vegetation-cover type use.
e 95% Bonferroni confidence interval constructed about the observed proportion
of vegetation-cover type use.
f Expected proportions of vegetation-cover type use less than the lower
confidence limit denote preference (P) (P<0.05), those exceeding the upper
confidence limit denote avoidance (A) (P<0.05), and those falling within the
confidence interval denote lack of selection (NS) (P>0.05).
9 Designation of vegetation-cover types: Gl_=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
64
Table 13. Proportional occupation of vegetation-cover types within home ranges
by adult female mule deer in relation to proportional availability of vegetationcover types within home ranges, Rosebud Mine / Colstrip, Montana, summer
1993-spring 1994.
Season3
Nb-
O0
Ed
Confidence interval6
Selectionf
SUM
GL9
MX
PS
RP
DS
RC
AG
UR
20
31
27
33
29
32
12
6
0.204
0.215
0.466
0.121
0.034
0.185
0.056
0.014
0.143 .
0.264
0.218
0.042
0.233
0.223
0.043
0.077
0.153
0.173
0.412
0.089
0.015
0.146
0.018
0.000
<
<
<
<
<
<
<
<
O
O
0
O
O
O
O
O
<
<
<
<
<
<
<
<
0,255
0.257
0.500
0.153
0.053
0.224
0.094
0.041
P
A
P
P
A
NS
NS
A
GL
MX
PS
RP
DS
RC
AG
UR
18
33
26
32
30
32
13
3
0.296
0.146
0.272
0.091
0.033
0.370
0.135
0.111
0.153
0.244
0.129
0.035
0.221
0.339
0.063
0.089
0.235
0.111
0.223
0.062
0.015
0.322
0.081
0.008
<
<
<
<
<
<
<
<
O <
0<
O <
O <
O <
O <
O <
O <
0.357
0.181
0.321
0.120
0:051
0.418
0.189
0.214
P
A
P
P
A
NS
P
NS
GL
MX
PS
RP
DS
RC
AG
UR
19
38
33
36
31
33
6
9
0.083
0.167
0.407
0.090
0.038
0.328
0.042
0.130
0.126
0.252
0.170
0.038
0.210
0.274
0.069
0.049
0.047
0.133
0.359
0.063
0.019
0.282
0.000
0.067
<
<
<
<
<
<
<
<
O <
0<
O <
O <
0<
O <
O <
O <
0.119
0.201
0.455
0.117
0.057
0.374
0.088
0.193
A
A
P
P
A
A
NS
P
GL
MX
PS
RP
DS
RC
AG
UR
20
32
27
33
29
28
13
5
0.238
0.190
0.340
0.078
0.049
0.258
0.141
0.167
0.198
0.241
0.171
0.034
0.241
0.210
0.067
0.053
0.184
0.151
0.288
0.052
0.026
0.211
0.086
0.073
<
<
<
<
<
<
<
<
O <
O <
O <
O <
O <
O <
0<
O <
0.292
0.229
0.392
0.104
0.072
0.305
0.196
0.261
NS
A
P
P
A
P
A
P
FAL
WNT
SPR
65
Table 13. Continued, 2 of 2.
a Seasonal designations: SUM=summer, 01 June-31 August; FAL=fall, 01
September-30 November; WNT=winter, 01 December-28 February; and
SPR=Spring, 01 March-31 May.
b The number of adult females from which selection of vegetation-cover types
was derived.
c The observed proportion of vegetation-cover type use.
d The expected proportion of vegetation-cover type use.
e 95% confidence interval constructed about the observed proportion of
vegetation-cover type use.
f Expected proportions of vegetation-cover type use less than the lower
confidence limit denote preference (P) (P<0.05), those exceeding the upper
confidence limit denote avoidance (A) (P<0.05), and those falling within the
confidence interval denote lack of selection (NS) (P>0.05).
9 Designation of vegetation-cover types: GL=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
66
Table 14. Proportional occupation of vegetation-cover types within home ranges
by adult male mule deer in relation to proportional availability of vegetation-cover
types within home ranges, Rosebud Mine / Colstrip1 Montana, summer 1993spring 1994.
Season8
Nb
O0
Ed
Confidence interval6
Selectionf
SUM
V
O
V
CD
O
d
0.127
0.349
0.014
0.000
0.097
0,013
0.000
<
<
<
<
<
<
<
O
O
O
O
O
O
O
<
<
<
<
<
<
<
0.147
0.289
0.547
0.112
0.030
0.261
0.153
0.000
NS
A
P
NS
A
NS
NS
A
0.069
0.304
0.090
0.032
0.211
0.269
0.022
0.002
0.000
0.138
0.126
0.003
0.000
0.303
0.000
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.105
0.334
0.318
0.109
0.000
0.531
0.099
0.000
NS
NS
P
NS
A
P
NS
A
0.000
0.104
0.604
0.167
0.000
0.104
0.000
0.082
0.093
0.330
0.098
0.059
0.246
0.127
0.026
0.020
0.000
0.018
0.466
0.061
0.000
0.018
0.000
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.000
0.190
0.742
0.273
0.000
0.190
0.000
0.239
A
A
P
P
A
NS
A
NS
0.063
0.313
0.417
0.021
0.021
0.083
0.333
0.000
0.105
0.244
0.304
0.034
0.088
0.207
0.011
0.007
0.000
0.182
0.278
0.000
0.000
0.005
0.066
0.000
<
<
<
<
<
<
<
<
O
O
O
O
O
O
O
O
<
<
<
<
<
<
<
<
0.132
0.444
0.556
0.062
0.062
0.161
0.600
0.000
NS
NS
NS
NS
A
A
P
A
GL9
MX
PS
RP
DS
RC
AG
UR
6
8
8
8
8
7
5
2
0.083
0.208
0.448
0.063
0.010
0.179
0.083
0.000
0.050
0.341
0.197
0.045
0.120
0.164
0.043
0.040
GL
MX
PS
RP
DS
RC
AG
UR
5
6
6
6
6
6
4
2
0.050
0.236
0.222
0.056
0.000
0.417
0.042
0.000
GL
MX
PS
RP
DS
RC
AG
UR
3
4
4
4
3
4
2
2
GL
MX
PS
RP
DS
RC
AG
UR
4
4
4
4
4
4
2
2
,
FAL
W NT
SPR
67
Table 14. Continued, 2 of 2.
a Seasonal designations: SUM=Summer, 01 June-31 August; FAL=fall, 01
September-30 November; WNT=winter, 01. December-28 February; and
SPR=spring, 01 March-31 May.
b The number of adult males from which selection of vegetation-cover types was
derived.
c The observed proportion of vegetation-cover type use.
d The expected proportion of vegetation-cover type use.
e 95% confidence interval constructed about the observed proportion of
vegetation-cover type use.
f Expected proportions of vegetation-cover type use less than the lower
confidence limit denote preference (P) (£<0.05), those exceeding the upper
confidence limit denote avoidance (A) (P<0.05), and those falling within the
confidence interval denote lack of selection (NS) (P>0.05).
9 Designation of vegetation-cover types: GL=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
(
68
1980, Ordway and Krausman 1986, Geist 1994a). Male use of pine savannah
types also was high during summer but use of riparian types was only moderate,
perhaps reflecting either less need for cover and / or succulent forage relative to
females, or the predominant use of vegetation-cover types with succulent forage
and cover by reproductive females. By fall, fawns either matured or died and the
predominant use within home ranges by females shifted from high-cover
vegetation-cover types such as pine savannah to relatively open reclamation
types. Males also made this shift, presumably to maximize breeding
opportunities. During winter, female use of reclamation and urban vegetationcover types increased while males tended to avoid reclamation and increased
use of riparian types. During spring, females again made extensive use of
reclamation and urban types as well as native grasslands while males avoided
these types.
Analyses involving use in relation to availability have been used widely to
infer preference (use greater than expected) or avoidance (use less than
expected) of specific vegetation-cover types (White and Garrott 1990).
Depending on the level of analysis, however, results can be misleading. For
example, adult females were broadly associated with disturbance types across
the study area (second-order preference), but these types rarely were used when
occurring within home ranges (third-order avoidance). Similar examples can be
cited for pine savannah and riparian types which generally were avoided or
received no selection at the second-order level, but always were preferred at the
third-order level. When results agree at both second- and third-order levels,
69
strong selection usually may be inferred. Even then, caution may be necessary in
interpretation of results. For example, mixed shrub types were avoided at both
levels of analysis by adult females, suggesting strong avoidance. Such a
conclusion, especially application of the term avoidance, is misleading, however,
because mixed shrub types comprised a very large portion of the study area and
most home ranges. Despite use less than availability, absolute use of mixed
shrub types was still quite high as indicated by observed proportions of
vegetation-cover type use (e.g., see Table 13).
Timing and nature of use by females varied substantially among individual
vegetation-cover types and subtly among seasons.
Females occupied “open”
types (grassland, reclamation, agriculture, and urban) primarily during nocturnal
hours (Table 15) and were predominantly active while doing so (Table 16). In
contrast, types providing concealment and thermal cover (pine savannah and
riparian) were occupied primarily during diurnal hours and used mostly as bed /
resting sites. These findings corroborate the general conclusion that deer forage
and rest principally during nocturnal and diurnal periods, respectively (Jacobson
1994).
Despite general trends, females refined the timing and nature of use of
specific vegetation-cover types to meet seasonal requirements for survival and
reproduction. For example, forbs were preferred diet items during summer, fall,
and spring (see food habits below) and occurred predominantly in open
vegetation-cover types. During these seasons, female use of vegetation and
cover was consistent with maximizing net energy gain, as they foraged
70
Table 15. Number of radiolocations obtained from adult female mule deer while
occupying each of 8 vegetation-cover types during diurnal and nocturnal periods,
Rosebud Mine / Colstrip, summer 1993-spring 1994.
Number of locations
Nocturnal
Diurnal
X2
Activity
status*
18
35
90 .
31
9
26
1
0
31
45
61
18
3
43
8
I
1.7
0.6
2.8
1.7
1.5
2.1
2.7
0.5
Equal
Equal
Equal
Equal
Equal
Equal
Equal
Equal
GL
MX
PS
RP
DS
RC
AG
UR
23
23
66
27
8
59
4
0
41
35
19
8
4
82
17
4
2.5
1.2
13.0
5.2
0.7
1.9
4.0
2.0
Equal
Equal
Diurnal
Diurnal
Equal
Equal
Nocturnal
Equal
GL
MX
PS
RP
DS
RC
AG
UR
7
47
77
26
5
63
0
3
12
29
84
13
9
67
3
11
0.7
2.1
0.2
2.2
0.6
0.1
1.5
2.3
Equal
Equal
Equal
Equal
Equal
Equal
Equal
Equal
GL
MX
PS
RP
DS
RC
AG
UR
18
46
74
18
12
30
4
2
39
30
36
13
6
54
18
8
3.9
1.7
6.6
0.4
1.0
3.4
4.5
1.8
Nocturnal
Equal
Diurnal
Equal
Equal
Nocturnal
Nocturnal
Equal
Season3
SUM
GLb
MX
PS
RP
DS
RC
AG
UR
FAL
.
WNT
SPR
71
Table 15. Continued 2 of 2.
Season3
Number of locations
Diurnal
Nocturnal
X *2
Activity
status*
113
139
200
52
22
246
46
24
6.2
0.2
11.3
8.1
1.3
5.5
12.4
6.2
Nocturnal
Equal
Diurnal
Diurnal
Equal
Nocturnal
Nocturnal
Nocturnal
ANN
GL
MX
PS
RP
DS
RC
AG
UR
66
151
307
102
34
178
9
5
* Denotes significance (Diurnal / Nocturnal) (P<0.05) or lack of significance
(Equal) (P>0.05).
3 Seasonal designations: SUM=summer, 01 June-31 August; FAL=fali, 01
September-30 November; WNT=winter, 01 December-28 February; and
SPR=spring, 01 March-31 May; annual, 01 June-31 May.
b Designation of vegetation-cover types: GL=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
72
Table 16. Number of radiolocations obtained from adult female mule deer while
active and inactive in each of 8 vegetation-cover types, Rosebud Mine I Colstrip,
summer 1993-spring 1994.
Number of locations
Active
Inactive
Season3
X2
Activity
status*
SUM
33
55
56
20
7
45
6
I
16
25
95
29
5
24
3
0
2.9
5.6
5.0
0.8
0.2
3.2
0.5
0.5
Equal
Active
Inactive
Equal
Equal
Equal
Equal
Equal
GL
MX
PS
RP
DS
RC
AG
UR
41
31
31
15
3 .
82
14
3
23
27
54
20
9
59
7
1
2.5
0.1
3.1
0.4
1.5
1.9
1.2
0.5
Equal
Equal
Equal
Equal
Equal
Equal
Equal
Equal
GL
MX
PS
RP
DS
RC
AG
UR
6
49
40
14
6
81
1
11
13
27
121
25
8
49
2
3
1.3
3.2
20.4
1.6
0.1
3.9
0.2
2.3
Equal
Equal
Inactive
Equal
Equal
Active
Equal
Equal
GL
MX
PS
RP
DS
RC
AG
UR
26
47
42
17
5
48
19
9
31
29
68
14
13
36
3
1
0.2
2.1
3.1
0.1
1.8
0.9
5.8
3.2
Equal
Equal
Equal
Equal
Equal
Equal
Active
Equal
GLb
MX
PS
RP
DS
RC
AG
UR
FAL
.
WNT
„
SPR
.
'
73
Table 16. Continued 2 of 2.
Number of locations
Active
Inactive
Season*3
X2
Activity
status*
ANN
GL
MX
PS
RP
DS
RC
AG
UR
106
182
108
66
21
256
40
24
83
108
169
88
35
168
15
5
1.4
9.4
6.7
1.6
1.8
9.1
5.7
6.2
Equal
Active
Inactive
Equal
Equal
Active
Active
Active
* Denotes significance (Active / Inactive) (P<0.05) or lack of significance (Equal)
(E>0.05).
3 Seasonal designations: SUM=summer, 01 June-31 August; FAL=fall, 01
September-30 November; WNT=winter, 01 December-28 February; and
SPR=spring, 01 March-31 May; annual, 01 June-31 May.
b Designation of vegetation-cover types: GL=grassland; MX=mixed shrub;
PS=pine savannah; RP=riparian; DS=disturbance; RC=reclamation;
AG=agriculture; and UR=urban.
74
nocturnally for preferred forbs among types lacking cover and bedded diurnally
among types providing cover. Lack of extreme nocturnal temperatures during
spring, summer, and fall presumably allowed deer to forage in cover-poor types
with minimal energy loss due to thermal conditions. During winter, availability of
forbs was reduced and browse became the principal diet item. Browse plants
occurred both in vegetation-cover types providing thermal cover such as riparian
and pine savannah and relatively open mixed shrub types.
Because of the
thermal constraints imposed by relatively low nocturnal temperatures during
winter, deer presumably benefited from the observed increase in nocturnal use
of pine savannah types. Benefits likely also were derived by the increased use of
relatively open mixed shrub types providing abundant browse during mild diurnal
hours.
Nocturnal Use of Colstrip
Nocturnal use of Colstrip by deer varied both spatially and temporally. The
mean city-wide UDI value was 0.79+1.13 (SE) (i.e. 1 deer observed every 1.26
km of census route) (Table 17). Annual UDI means differed among sections of
Colstrip (X2=17.395, df=4, P=O.002). Section B received the greatest use,
followed by Sections A, D, C, and E in decreasing order, with each sectional
mean differing from all others (P<0.05). UDI means also differed among months
(X2=78.392, D F=11, P=0.000) and were highest and lowest during fall / winter
and spring / summer months, respectively. Each monthly mean differed from all
others (P<0.05) with the exception of February and July (P>0.05).
75
Table 17. Mean (±SE) Urban Deer Index (UDI) values for mule deer among
sections of Colstrip, Montana, July 1993-June 1994.
A
Month
July
August
September
October
November
December
January
February
March
April
May
June
Annual
B
Section of Colstrio
D
C
±0.00
0.00
±0.00
0.30
±0.52
2.79
±1.64
2.33
±1.19
1.88
±1.45
0.53
±0.91
0.00
±0.00
2.18
±0.69
0.30
±0.52
0.00
±0.00
0.15
±0.26
0.15
±0.17
0.51
±0.31
0.46
±0.47
1.03
±0.44
2.32
±1.07
3.34
±0.76
3.14
±2.02
0.17
±0.81
1.44
±1.48
1.34
±1.28
2.26
±1.08
0.21
±0.21
0.00
±0.00
0.30
±0.18
0.03
±0.05
1.57
±0.92
1.46
±0.64
1.21
±0.93
0.44
±0.28
0.08
±0.09
1.30
±1.11
0.64
±0.44
0.50
±0.50
0.00
±0.00
0.27
±0.23
0.08
±0.08
0.31
±0.33
1.82
±1.30
1.74
±0.61
0.08
±0.13
0.85
±0.32
0.43
±0.74
1.51
±0.84
1.66
±1.47
0.23
±0.40
0.08
±0.13
0.87
±1.32
1.41
±1.46
0.63
±0.81
0.75
±0.98
0 .0 0 a
E City-wide
.
0.23
±0.23
0.52
±0.30
0.12
±0.12
0.58
±0.48
0.52
±0.34
0.70
±0.64
0.70
±0.77
0.00
±0.00
0.00
±0.00
1.28
±0.94
0.12
±0.20
0.00
±0.00
0.13
±0.07
0,29
±0.08
0.21
±0.09
1.53
±0.59
1.63
±0.20
1.32
±0.40
1.05
±0.40
0.24
±0.30
1.28
±0.47
1.05
±0.57
0.63
. ±0.19
0.07
±0.05
0.40
±0.59
0.79
±1.13
a Mean Urban Deer Index (UDI) values were calculated by dividing the number of
deer observed during 4 monthly nocturnal surveys in a given section of Colstrip
by the number of kilometers driven during the course of the survey in that
section.
Nocturnal use of Colstrip was principally by deer with home ranges
centered near the townsite. The greatest known distance traveled by a
radiocollared deer during the course of 1 day from a diurnal location outside of
/
76
Colstrip to a nocturnal location within Colstrip was 1734.7 m, but the mean was
considerably less (656.2+498.3 (SE) m, n=36 paired observations).
Residence and landscaping characteristics varied greatly among the 5
sections of Cplstrip. City-wide, houses and mobile homes predominated, while
apartments were relatively uncommon (Table 18). Houses were most common in
Sections B and C and mobile homes in Sections A, D, and E. Most residences
city-wide (71.3% ) contained landscape plantings, 21.4% contained plantings that
were protected from potential deer browsing, and 4.6% contained plantings that
exhibited obvious signs of deer browsing (Table 19). These trends were fairly
uniform among sections of Colstrip, although landscape plantings were more
likely present in yards of houses than mobile homes (2=8.025, P=0.000), and
houses were most abundant in Sections B and C.
Table 18. Percentage occurrence of houses, mobile homes, and apartments
among sections of Colstrip, Montana, January 1994.
Residence type
House
Mobil home
Apartment
A
18.8
63.6
17.6
Section of Colstrio
C
B
76.1
0.0
23.9
90,0
0.0
10.0
D
E
City-wide
5.9
94.1
0.0
24.2
75.8
0.0
47.2
41,6
10.7
o
77
Table 19. Percentage occurrence of residences containing landscape plantings,
protected plantings, and deer-damaged plantings in Colstrip, Montana, January
1994.
Plantings
A
B
Present
Protected
Damaged
55.9
20.0
4.7
92.2
29.9
7.5
Section of Colstrip
D
C
88.9
22.7
5.5
54.4
11.8
2.9
E
City-wide
61.3
22.5
1.6
71.3
21.4
4.6
Deer use of Colstrip was influenced by inherent deer behaviors and
seasonal variation of habitat conditions. For example, deer presence in Colstrip
was very low during late spring and summer. Female avoidance of Colstrip
during this time may be explained by the tendency for females to occupy, when
possible, reproductive habitats that best provide for the requirements of female
lactation and fawn survival (Pac et al. 1991). Speculatively, females may have
avoided Colstrip during fawning-rearing because urban areas lacked either
adequate forage resources to support female survival and lactation or adequate
isolation or concealment cover to enhance fawn survival. Absence of males from
Colstrip during summer may be explained by the ability of natural vegetationcover types to adequately provide for survival and growth requirements of males
(Cook 1972, Parker 1994), thereby rendering use of urban areas unnecessary.
Deer use of Colstrip increased markedly during fall and winter, specifically
after the first hard freeze and snow accumulation on 08 October 1993. At this
time, remaining succulent forbs outside of Colstrip likely (1) dried and became
less palatable due to physiological responses to the freeze (Short et al. 1972) and
I
78
(2) became less available due to the accumulation of snow. At the same time,
forage resources (fallen crabapples) became available in Colstrip, and deer with
home ranges peripheral to the townsite made greater use of urban areas.
Crabapples were abundant in Section A of Colstrip and available to a lesser
degree in Sections B and C, and deer frequently were observed consuming them.
Accordingly, UDI values were highest in Section A during October and
November, the months during which availability of crabapples was greatest. By
the end of November, however, deer distribution shifted to Sections B and C,
presumably because (1) the availability of crabapples decreased in Section A due
to cumulative use and increasing snow depth, and (2) alternative food resources
(palatable landscape plantings) were available in Sections B and C. Deer use in
these sections remained high throughout winter and early spring, with the
exception of February. Deer abundance in Section A peaked a second time after
snow melt in March made previously covered crabapples available once again.
Extreme winter weather conditions apparently impacted deer use of
Colstrip during February, the coldest month of the study period, and the month
during which snow depth was consistently greatest. When necessary, deer
conserve energy during times of extreme winter weather by reducing mobility
(Mackie 1994a) and selecting habitats that provide protection from radiant and
convective heat loss (Mackie 1994b). During February, radiocollared deer
generally reduced mobility and activity, and either increased use of wooded
portions of their home ranges or moved to winter accessory areas providing
greater thermal cover. These observations suggested that (1) under weather
/
79
conditions that prevailed during February, deer exhibited behavior consistent with
energy conservation, and (2) this strategy was incompatible with nocturnal
foraging in Colstrip.
Food Habits
Food habits of mule deer differed seasonally and generally confirmed
findings from other studies (Wilkins 1957, Kamps 1969, Mackie 1970, Kufeld et
al. 1973, Schwarzkoph 1973, Hamlin 1974, Knowles 1975, Komberec 1976,
Nyberg 1980, Pac et al. 1991). Seasonal use of grasses and forbs by deer on
the Rosebud Mine / Colstrip study area was somewhat lower and higher,
respectively, than expected based on past research (Table 20). Shrubs were the
dominant food item during winter but also comprised a large percentage of spring
and fall diets. Forbs dominated during spring, summer, and fall, forming >50% of
the diet during each season. Grasses, although never a principal component,
occurred most often in diets during spring and fall.
Numerous diet items were identified to the species level. Forbs found in
rumens most consistently across all seasons included alfalfa, yellow sweetclover,
and common salsify (Traaopoaon dubius). Commonly encountered shrubs
included silver sagebrush, fourwing saltbush, fragrant sumac, and common
snowberry. The relatively high percentage occurrence Of forbs and low
occurrence of grasses and shrubs in seasonal diets likely resulted from the
abundance of 2 preferred food items, alfalfa and yellow sweetclover, among most
vegetation-cover types. Both forbs were distributed widely throughout the study
80
Table 20. Mean percentage composition of diet items, by major forage class, and
frequency occurrence (%) of individual plant species contained within rumens
obtained from female mule deer, Rosebud Mine / Colstrip, Montana, fall 1992summer 1994.
Forage Classb
Speciesc
Grasses
Aaropvron cristatum
Avena fatua
Bromus tectorum
Forbs
Ambrosia psiIostachva
Artemisia friaida
Astragalus spp.
Brassica spp.
Cirsium undulatum
Convolvulus arvensis
Gutierrezia sarothrae
Kochia scoparia
Lactuca serriola
Linum perenne
Malva parviflora
Medcaao sativa
Melilotus officinalis
Ratibida columnifera
Salsola kali
Taraxacum officinale
Traaopoaon dubius
Verbena thapsus
Yucca alauca
Shrubs
Artemisia cana
Artemisia tridentata
Atriplex canescens
Chrvsothamnus nauseosus
Elaeaanus anaustifolia
Juniperus scopulorum
Pvrus spp.
Pinus ponderosa
Prunus americana
Prunus virainiana
Rhus aromatica
Rosa spp.
SUM
Season3
WNT
FAL
SPR
3.9
7.7
0.0
0.0
88.2
0.0
7.7
38.5
0,0
0.0
30.8
0.0
15.4
15.4
23.1
2.2
69.2
61.5
7.7
0.0
7.7
46.2
7.7
0.0
7.9
30.8
0.0
23.1
7.7
15.4
0.0
0.0
0.0
0.0
0.0
23.1
7.7
6.4
0.0
4.3
6.5
50.5
0.0
10.9
8.7
10.9
4.3
6.5
2.2
13.0
6 .0
2.2
0.0
52.2
43.5
0.0
2.2
2.2
17.4
0.0
8.7
42.8
41.3
2.2
43.5
0.0
2.2
6.5
2.2
4.3
0.0
4.3
23.9
4.3
4.0
0.0
0.0
0.0
16.1
0.0
0.0
0.0
0.0
7.1
0.0
0.0
0.0
7.1
0.0
0.0
64.3
14.3
0.0
0.0
0.0
28.6
7.1
35.7
79.4
35.7
7.1
50.0
7.1
14.3
42.9
14.3
28.6
0.0
7.1
28.6
0.0
8.3
5.9
0.0
29.4
55.9
2.2
5.9
5.9
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
76.5
35.3
5.9
0.0
0.0
29.4
0.0
0.0
35.6
35.3
11.8
11.8
5.9
0.0
5.9
0.0
0.0
5.9
0.0
29.4
0.0
,
.
81
Table 20. Continued, 2 of 2.
Forage class'3
Species0
SUM
Salix spp.
Svmphoricarpos albus
7.7
15.4
Season3
FAL
WNT
SPR
2.2
54.3
0.0
41.2
0.0
57.1
a Seasonal designations: SUM=summer, 01 June-31 August; FAL=fall, 01
September-30 November; WNT=winter, 01 December-28 February; and
SPR=spring, 01 March-31 May.
b Seasonal use for the 3 major forage classes (grasses, forbs, and shrubs /
trees) is given as the mean percentage composition, as determined by pointframe estimation, of 13 rumens obtained during summer, 46 during fall, 14
during winter, and 17 during spring.
c Seasonal use for individual plant species is given as the percentage occurrence
of each item in all rumens obtained during a given season.
r
82
area due to agricultural and reclamation practices as well as favorable summer /
fall precipitation patterns during the study. The relatively high use of forbs,
particularly during fall and spring, confirms that forage availability was high during
the study, and deer did not have to resort to consuming woody browse of lower
nutritional value,
Physical Condition .
Physical measurements obtained from 93 adult females indicated condition
Of deer was highest and lowest during fall and spring, respectively (Table 21).
Both whole (X2= 1 1.324, DF=3, £=0.010) and dressed (X2=23.905, DF=3,
£=0.000) weights differed seasonally, fall weights exceeding those during
summer, winter, and spring (P<0.05). Measures of kidney-fat also differed
seasonally (X2=42.180, DF=3, P=0.000), fall index values exceeding those during
spring (P<0.05). Similar patterns of weight gain / loss and kidney-fat deposition /
depletion were exhibited by yearling and adult females. Deer inhabiting adequate
environments accumulate body fat and increase condition during summer and
autumn, exploiting abundant, high-quality forage, and lose fat and decrease
condition during winter and spring as forage availability and nutritional quality
decrease and energetic demands of survival and reproduction increase
(Anderson and Bowden 1972, Cook 1972, Wallmo 1978, Short 1981, Brown
1994).
83
Table 21. Mean (±SE) whole and dressed weights and kidney fat index values
for adult female mule deer, Rosebud Mine / Colstrip, Montana, fall 1992-summer
1994.
Season®
Paramterb
Whole weight (kg)
Dressed weight (kg)
Kidney-fat index
SUM
FAL
WNT
SPR
66.2
±24.8
44.2
±5.2
0.2
±0.2
68.1
±6.4
49.7
±5.5
1.3
±0.7
62.6
±4.9
43.7
±4.7
0.6
±0.2
59.3
±9.6
39.0
±13.1
0.2
±0.2
a Seasonal designations: SUM=summer, 01 June-31 August; FAL=fall, 01
September-30 November; WNT=winter, 01 December-28 February; and
SPR=spring, 01 March-31 May.
b Summer, fall, winter, and spring estimates were based on examination of 13,
51, 13, and 16 adult female mule deer, respectively.
Population Characteristics
Abundance and Trend
Population estimates varied considerably among seasonal survey flights
but indicated a relatively high density of mule deer for a prairie-environment
population (Mackie 1994c). Estimates obtained during fixed-wing (n=9) and
helicopter (n=4) flights averaged 1,714±1068 (SE) deer (7.2 deer I km2) and
1,800±304 (SE) deer (7.6 deer / km2), respectively (Table 22).
More deer always
were observed during winter and spring helicopter flights (m ean=1109+104 (SE)
deer) than during fixed-wing flights (mean=433±149 (SE) deer) (t=5.180, DF=6,
P=0.002). Further, sightability of radiocollared deer was greater during helicopter
(mean=64.4% of available radiocollared deer sighted per survey, range=47.892.0%) than fixed-wing flights (mean=31.8% of available radiocollared deer
84
Table 22. Lincoln index population estimates for mule deer, Rosebud Mine /
Colstrip, Montana, summer 1992-spring 1995.
Flight3
FXW
FXW
FXW
FXW
FXW
FXW
FXW
FXW
FXW
Seasonb
SUM
FAL
WNT
SPR
SUM
FAL
WNT
SPR
SUM
FXW
HLC
HLC
HLC
HLC
HLC
WNT
SPR
WNT
SPR
Year
nc
Md
me
Nf
Confidence interval9
1992
1992
1992
1993
1993
1993
1993
1994
1994
367
556
473
748
318
520
225
329
362
16
16
16
52
49
49
43
40
40
3
2
11
24
11
22
11
12
7
1957
4448
688
1620
1416
1158
879
1096
2172
0<
0<
277 <
969 <
576 <
679 <
361 <
474 <
619 <
Mean
433
35
11
1377
556 < N < 2 1 9 7
1993
1994
1994
1995
1099
1111
1261
966
43
42
25
23
22 2148
28 1666
23 1370
11 2019
Mean
1109
33
21
1742
1241 <
1044<
803 <
807 <
N
N
N
N
N
N
N
N
N
N
N
N
N
<4167
< 9727
< 1098
< 2270
< 2255
< 1636
< 1396
< 1717
< 3724
< 3054
<2287
< 1936
< 3230
989 < N < 2495
a Aerial survey flights from which estimates were derived were conducted either
with a fixed-wing aircraft (FXW ) or helicopter (HLC).
b Seasonal designations: SUM=summer, 01 June-31 August; FAL=fall, 01
September-30 November; WNT=winter, 01 December-28 August; and
SPR=spring, 01 March-31 May.
0 The number of deer observed during an aerial survey.
d The number of marked deer potentially observed during an aerial survey.
6 The number of marked deer observed during an aerial survey.
f The Lincoln index population estimate.
9 The 95% confidence interval constructed around the Lincoln index value.
sighted per survey, range=12.5-68.8% ) (t=2.903, DF=11, P=0.014). Estimates of
deer abundance did not differ between flight types (t=0.145, D F=11, P=0.887),
although estimates obtained from helicopter flights varied less seasonally and
had smaller confidence intervals.
Trends in deer abundance since 1974, indexed by the number of animals
85
observed per survey hour, generally decreased 1974-1980, increased 1981-1989,
and remained relatively stable 1990-1994 (Figure 10).
Trends followed those
exhibited by other mule deer populations in eastern Montana (Mackie et al. 1985,
Hamlin and Mackie 1989, Wood et al. 1989), although timing of population lows
and subsequent recovery apparently lagged by 2-3 years. This possibly resulted
from more restricted hunting in the Colstrip area. The observed increase in deer
numbers through time roughly paralleled increased mining development in the
area. Phillips et al. (1985) reported a similar correlation on a surface coal mine in
southeastern Montana and attributed it in part to the beneficial modification of
topography and vegetation resulting from mining-related disturbance. Although
mining-related alteration of habitat may be partially responsible for increasing
abundance of deer over the last 20 years, mule deer populations throughout the
western U S. were low when collection of population data on the Rosebud Mine /
Colstrip study area began (Julander and Low 1976). Accordingly, even modest
recovery from population lows during the mid-1970s coincident with mining
development could falsely suggest a more positive relationship between surface
mining and mule deer population growth than actually existed.
Distribution of deer shifted from 1974 to 1993 as deer numbers in areas
close to Colstrip and surrounding reclamation increased faster than numbers of
deer in outlying areas (Appendix, Figures 13-32). Mean distance from the center
of Colstrip to each deer group observed during aerial surveys was 7.0+3.3,
6.4±3.9, and 5.8±4.1 (SE) km during the periods 1974-1981 (deer population
decreasing), 1982-1989 (deer population increasing), and 1990-1993 (deer
86
o
70
2
40
Q
20
1974
1976
1978
1980
1982
1984
1986
1988
1990
1992
199/
Year
Figure 10. Mean number of mule deer observed per hour during aerial surveys of
the Rosebud Mine permit area, 1974-1994. Means from 1974-1982 were derived
from monthly surveys (n=12) flying 1-mile transects. Means from 1983-1985
were derived from seasonal, full-coverage surveys (n=4). Means from 1986-1994
were derived from seasonal surveys (n=4) flying 0.5-mile transects.
population stable), respectively. Values differed among time periods (X2=75.024,
DF=2, P=0.000), with each mean differing from the others (P<0.05). That deer
distribution shifted toward Colstrip as the population increased and stabilized
suggests one of two possibilities: (1) that core reproductive habitats were located
in areas distant from the townsite and adjacent mining / reclamation areas, and
as deer numbers increased, individuals increasingly colonized and occupied more
marginal habitats amidst mining and urban development; or (2) as mining
development increased 1974 to present, habitat improvements allowed
87
abundance to increase in mining and urban areas as once marginal habitats
became more conducive to successful reproduction by deer. The second
alternative seems more likely, as the mean number of observed fawns per grid
cell (all aerial surveys 1992-1994) was greater for cells containing reclamation
than for those lacking it (sign rank =18.0, P=0.008).
Aae / Sex Structure
The population was characterized by moderate to low survival of fawns
and low survival of males during 1992-1995. Numbers of fawns: 100 females and
'i
males: 100 females in winter and spring were relatively consistent between years,
averaging 50.9+3.7 and 28.9±0.6 (SE), respectively (Table 23). Fawn:doe ratios
were low relative to estimates obtained for other eastern Montana populations
(Hamlin and Mackie 1989, Jackson 1990).. They agreed more closely with ratios
for mountain-foothills populations in southwestern Montana (Pac et al. 1991),
possibly suggesting a population with relatively low, stable rates of annual
turnover and fawn recruitment (Mackie 1983).
A mean adult sex ratio of 28.9 males: 100 females during early winters
1993 and 1994 was unexpectedly low given that males were not hunted on the
area. Wood et al. (1989) and Hamlin and Mackie (1989) reported autumn
male:female ratios of 35:100 and 42.2:100, respectively, for hunted populations.
However, Martinka (1978), Gavin et al. (1984), and Kie and White (1985)
reported sex ratios similar to this study for deer populations in which males were
not harvested. These findings support conclusions that females live longer than
88
Table 23. Mule deer population density estimates (deer / km2) and relative
occurrence of fawns, adult females, and adult males derived from aerial surveys
(full-coverage helicopter flights only), Rosebud Mine / Colstrip, Montana, winter
1993-spring 1995.
Season3
WNT
SPR
WNT
SPR
Mean±SE
Year '
1993
1994
1994
1995
Fawns: 100
females
Fawns: 100
adults
Males: 100
females
9.1
7.1
5.8
8.6
47.2
NA
54.6
NA
36.4
36.2
42.6
31.4
29.5
NAc
28.3
NA
7.7±1.3
50.9+3.7
36.7±4.0
Density"
28.9±0.6
a Seasonal designations: WNT=winter, 01 December-28 February; and
SPR=Spring, 01 March-31 May.
b Density estimates (deer / km2) were derived from Lincoln estimates (see Table
22 ).
0 Estimate not available because sex of deer was not determined during spring
flights.
males regardless of harvest pressures. This could reflect physiological stresses
on males associated with rut,, habitat-selection patterns that predispose males to
higher rates of natural predation, or yearling dispersal and rut-related movements
that expose males to increased risk of mortality (Taber and Dasmann 1954,
Mackie 1994d). Rut-related movements led at least some males on the Rosebud
Mine into areas subject to hunting. This resulted in unexpectedly high mortality
among males and contributed to the observed male:female ratio.
Age structure, based on hunter harvested, vehicle killed, and collected
deer, was heavily skewed toward young age classes for males and toward older
age classes for females (Figure 11). The mean age for females and males was
5.7±4.2 and 2.4±2.6 (SE) years, respectively. Among males, 64.0% were <2.5
<
89
22
20
18
0.5
1.5
2.5
3.5
4.5
5.5
6.5
Age (years)
Female
7.5
8.5
9.5
10.5+
Male
Figure 11. Age-specific numbers of female and male mule deer obtained from
hunter harvest, deer-vehicle collision, and special collection, Rosebud Mine /
Colstrip, Montana, September 1992-June 1994.
years of age. Only 21.3% of all females were <2.5 years of age. Further, 18.5%
of all females examined were >10.5 years of age. These findings reflect much
greater annual turnover or dispersal from the area among males as compared to
females. Again, this suggests a population more stable than typical for
populations inhabiting highly variable prairie environments.
Social Organization
Mean numbers and sizes of female, male, and mixed-sex groups observed
during ground surveys on the Rosebud Mine reflected seasonal changes in deer
90
sociality imposed by reproductive and survival behaviors (Table 24). Overall,
mean group size peaked during winter / early spring when males and females (at
least females without fawns) were aggregated on common wintering grounds.
Group size was lowest during summer when deer were most dispersed. Maleonly groups were observed most commonly and achieved their largest size during
the summer and early fall when segregation of the sexes generally is greatest
(Main 1994b). Thereafter, the number and size of male-only groups decreased,
and the number and size of mixed-sex groups increased, coincident with the
onset of rut. The largest number of female-only groups and those of the smallest
size were observed during summer when reproductive females exhibited typical
isolative and non-sociable behavior resulting in small, highly dispersed groups of
single adult females and their young of the year (Geist 1990 and 1994a, Main
1994b). The number of female-only groups decreased from summer to fall and
winter as cumulative fawn mortality released increasing numbers of females from
habitat-use constraints associated with fawn-rearing, resulting in reformation of
matrilineal groups (Hamlin and Mackie 1989).
Conception and Birth Dates
Conception and parturition dates exhibited by females during this study
closely matched those reported for other eastern Montana deer populations
(Hamlin and Mackie 1989, Jackson 1990). The median conception date for
yearling / adult females was 20 November based on fetal measurements, and
95% of all conceptions occurred 09-29 November based on a mean breeding
91
Table 24. Mean sizes and numbers of male, female, and mixed-sex mule deer
groups observed during ground surveys, Rosebud Mine / Colstrip, Montana, June
1993-May 1994.
6
7
8
9
10
1.9
1.4
1.6
3.8
2.0
1.8
1.7
3.5
2.3
1.6
2.3
3.3
3.3
1.6
3.5
5.0
5.2
1.3
4.5
9.6
4.8
1.0
4.9
5.9
3.8 3.8
8.0 11.0
2.3 2.3
2.3
8.5
1.0
1.3 0.8
5.5 10.3
0.8 1.8
0.8
6.3
3.5
Parameter®
Group size
All groups
Male groups
Female groups
Mixed-sex groups
Number of groups
Male groups
Female groups
Mixed-sex groups
Month
11
12
1
2
3
4
5
4.6
1.0
4.5
5.8
7.5
1.5
7.6
4.0
7.3
1.5
6.7
8.2
9.3
NA
NA
NA
7.3
NA
NA
NA
4.5
NA
NA
NA
0.8
3.5
1.5
0.5
2.3
0.3
0.0
1.5
1.0
NA
NA
NA
NA
NA
NA
NA
NA
NA
a Mean group number and size values were derived from 4 ground surveys
conducted along a survey route running through the Rosebud Mine. Deer sex
was not determined during March, April, and May surveys.
' <
date of 19 November±5 (SE) days. An independent estimate of the breeding
period was derived from capture dates of neonate fawns (n=31, median capture
date=08 June, mean capture date=09 June±5 (SE) days). Back-calculation from
the median capture date, assuming a gestation length of 203 days (Robinette et
al. 1973), resulted in a median conception date of 17 November; 95% of all
breeding occurred 07-27 November. The back-calculation method likely yielded a
slightly early estimation of conception date because capture of neonate fawns
began when the first fawn was observed and ended when all available
radiocollars were distributed. Doubtless, some fawns were born after capture
operations ceased.
92
Ovulation and Fertilization Rates
Ovulation rate, derived from the ovaries of 45 female reproductive tracts
collected December-June, was 1:68+0.44 (SE) ova per yearling / adult female;
none of 5 female fawns examined exhibited signs of breeding. As expected
(Robinette et al. 1957, Connolly 1981, Wood 1986, Sadleir 1987), the ovulation
rate of adults (n=31, 1.81 ±0.31 (SE) ova per female) exceeded that of yearlings
(n=14, 1.20+0.43 (SE) ova per female) (X2=7.674, DF= 1, P =0.006). Fertilization
rate for ova was 100.0% for both yearlings and adults. Findings from this study
were similar to those obtained elsewhere in eastern Montana (Eustace 1971,
Hamlin and Mackie 1989) and generally reflected adequate nutrition and high
productivity of females (Verme 1969). Thus, moderate to low fawmdoe and
fawn:adult ratios observed on the study area reflected high fawn mortality rather
than low basic productivity.
Secondary Sex Ratios
The combined sex ratio for collected fetuses (n=52) and captured neonate
fawns (n=41) was skewed significantly in favor of males (33 females:60 males)
(X2=3.919, D F = I, P=0.048). Analysis of the 2 groups individually, however,
indicated females and males occurred equally among fetuses (22 females:30
males, X2=0.615, DF=1, P=0.408), but females were less prevalent than males
among captured neonate fawns (11 females:30 males, X 2=4.402, DF=1,
P=0.038). Although relationships were not significant (P>0.05), yearling females
produced equal numbers of females and males (n=10 fetuses, 5 female and 5
93
male), prime-aged adult females (2.5-9.5 years old) produced half as many
females as males (n=36 fetuses, 12 female and 24 male), and old-aged adult
females (10.5 years and older) produced 5 times as many females as males (n=6
fetuses, 5 female and 1 male). Paired comparisons of male / female weights
indicated males were larger than females when both were produced in a single
litter (sign rank=30.0, DF=11, P=0.016).
The adaptive significance and causal mechanisms regarding adjustment of
secondary sex ratios among deer have been discussed at length (McCullough
1979, Verme and Ozoga 1981, Ozoga and Verme 1982, Verme 1983, Degayner
and Jordan 1987, Jacobson 1994). Although the overall sex ratio of fetuses and
neonate fawns on the Rosebud Mine / Colstrip study area was skewed toward
males, finer-scale analysis indicated no such relationship for fetuses alone. The
overall relationship was influenced primarily by the sex ratio of neonate fawns,
suggesting differential post-natal mortality of the sexes. The most plausible
explanation for differential mortality relates to fawn condition at birth. In this study
as well as others (Robinette et al. 1973, Clutton-Bfock et al. 1982, Putman 1988),
males were heavier at birth, or shortly thereafter, than their female siblings,
indicating greater pre-natal investment in male than female offspring. Further,
this relationship tends to be maintained during fawn-rearing as males nurse
longer and grow faster than female siblings (Blaxter et al. 1974, Glutton-Brock et
al. 1982). Thus, females are born smaller and grow slower than males, likely
resulting in comparatively high in utero death, abandonment at birth, or predation
shortly after birth, all of which would result in fewer female fawns available for
94
capture and, thus, a male-biased sex ratio.
Fawn Survivorship
Survivorship of radiocollared fawns (both sexes combined) was 0.336 and
0.545 during 1993-1994 and 1994-1995, respectively (Appendix, Tables 32 and
33). Log-rank tests indicated survivorship did not differ between years (P>0.05);
thus, data were pooled yielding an overall annual survivorship estimate of 0.431
(Table 25). Sixteen of 19 fawn deaths (84.2% ) occurred during either summer
(n=8) or winter (n=8) (Table 26). Coyote predation was the major source of
mortality, accounting for 14 of 19 deaths (Table 26). Although coyotes most often
were the proximal agents of mortality during winter and spring, poor body
condition resultant from winter stress and malnutrition likely predisposed most
fawns to predation. Poor condition was known to be the ultimate cause of death
for 1 fawn killed by coyotes during March 1994. Vehicle-related accidents, both
on and off payed roads, accounted for 10.5% of fawn deaths. Although crossing
roads accounted for the death of 1 fawn, another was killed by mining equipment
while bedded well away from the nearest road. With the relatively high volume of
road traffic and mining activity that occurs around the clock on a daily basis,
vehicle-related accidents may constitute a significant factor of mortality for fawns
on the study area.
Adult Survivorship
Annual survivorship of radiocollared adult females was high, ranging from
0.868 to 0.947, 1992-1995 (Appendix, Tables 34-36). Log-rank tests indicated
95
Table 25. Monthly survival estimates and 95% confidence intervals for
radiocollared female and male mule deer fawns, Rosebud Mine / Cdlstrip,
Montana, June 1992-May 1995.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk
35
31
27
25
24
22
21
20
15
12
11
10
#
#
deaths censored
4
2
2
0
1
0
0
4
3
1
0
0
0
2
0
1
1
1
1
1
0
0
1
1
#
added
0
0
0
0
0
0
0
0
0
0
0
0
Survival
0.886 .
0.829
0.767
0.767
0.735
0.735
0.735
0.588
0.471
0.431
0.431
0.431
. Confidence interval
0.787 < S <
0.708 < S <
0.628 < S <
0.622 < S <
0.584 < S <
0.577 < S <
0.573 < S <
0.423 < S <
0.297 < S <
0.247<S<
0.239 < S <
0.230 < S <
0.985
0.949
0.907
0.912
0.887
0.893
0.897
0.754
0.644
0.615
0.624
0.630
96
Table 26. Seasonal causes of death for radiocollared female and male mule deer
fawns, Rosebud Mine / Colstrip, Montana, June 1993-May 1995.
Deer ID
C076-93
C082-93
C085-93
C088-93
C090-93
C092-93
C095-94
C100-94
0073-93
0030-92
0074-93
0079-93
0086-93
0096-94
0097-94
0099-94
0101-94
0031-92
0077-93
Age at death
8 days
14 days
20 days
3 months
1 month
1 month
3 months
12 days
4 months (
7 months
7 months
8 months
8 months
9 months
8 months
8 months
9 months
9 months
10 months
Date of death
13 JUN
19 JUN
.3 0 JUN
27 AUG
11 JLY
23 JLY
28 AUG
22 JUN
01 O C T
15 JAN
07 JAN
21 JAN
01 FEB
18 FEB
26 JAN
26 JAN
18 FEB
15 JAN
29 MAR
1993
1993
1993
1993
1993
1993
1994
1994
1993
1993
1994
1994
1994
1995
1995
1995
1995
1993
1994
Season3
Cause of death
Summer
Summer
Summer
Summer
Summer
Summer
Summer
Summer
Fall
Winter
Winter
Winter
Winter
Winter
Winter
Winter
Winter
Spring
Spring
Coyote predation
Coyote predation
Coyote predation
Unknown
Coyote predation
Vehicle collision
Vehicle collision
Coyote predation
Coyote predation
Unknown
Coyote predation
Coyote predation
Coyote predation
Coyote predation
Coyote predation
Coyote predation
Coyote predation
Unknown
Coyote predation
a Seasonal designations: summer, 01 June-31 August; fall, 01 September-30
November; winter, 01 December-28 February; and spring, 01 March-31 May.
survivorship did not differ among years (P>0.05); thus, data were pooled yielding
an overall annual survivorship estimate of 0.900 (Table 27). As with fawns, the 2
known major causes of death for adult females were coyote predation and
vehicle-related accidents, which collectively accounted for 6 of 10 deaths (Table
28). Predation was distributed equally throughout the seasons, but vehiclerelated accidents occurred only during spring, perhaps a result of increased
mobility and home range expansion by females as vegetative growth began.
97
Table 27. Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult female mule deer, Rosebud Mine / Colstrip,
Montana, June 1992-May 1995.
#
at risk
Month
#
#
deaths censored
87
92
91
90
89
89
82
77
104
104
100
97
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
0
0
0
1
0
2
I
0
0
3
3
0
0
2
1
0
0
5
5
1
0
I
0
3
#
added
Survival
5
1
0
0
0
0
1
28
0
0
0
0
1.000
1.000
1.000
0.989
0.989
0.967
0.955
0.955
0.955
0.927
0.900
0.900
Confidence interval
1.000 < S <
1.000 < S <
1.000 < S <
0.967 < S <
0.967<S<
0.930 < S <
0.911 < S <
0.910 < S <
0.916 < S <
0.879 < S <
0.844 < S <
0.843 < S <
1.000
1.000
1.000
1.000
1.000
1.000
0.999
1.000
0.994
0.975
0.955
0.956
Table 28. Seasonal causes of death for radiocollared yearling / adult female
mule deer, Rosebud Mine / Colstrip, Montana, June 1992-May 1995.
Deer ID
C007-92
C 0 12-92
C047-93
C072-93
C003-92
C 0 14-92
C034-93
C038-93
C060-93
C066-93
Age at death
4.5
4.5
3.5
4.5
9.5
4.5
9.5
1.5
9.5
8.5
years
years
years
years
years
years
years
years
years
years
Date of death
15 SPT
15 NOV
26 NOV
15 DEC
01 APR
15 MAR
13 APR
13 APR
14 MAR
01 MAR
1992
1994
1994
1994
1994
1993
1995
1993
1994
1993
Season3
Cause of death
Fall
Fall
Fall
Winter
Spring
Spring
Spring
Spring
Spring
Spring
Coyote predation
Hunter harvest
Unknown
Coyote predation
Malnutrition
Vehicle collision
Vehicle collision
Vehicle collision
Coyote predation
Unknown
a Seasonal designations:, summer, 01 June-31 August; fall, 01 September-30
November; winter, 01 December-28 February; and spring, 01 March-31 May.
Annual survivorship of radiocollared males ranged from 0.500 to 0.909,
1992-1995 (Appendix, Tables 37-39). Log-rank tests again indicated no
98
differences among annual survivorship patterns (P>0.05), so data were pooled
yielding an overall annual survivorship estimate of 0.577 (Table 29). Hunter
harvest was the only known cause of mortality and was responsible for 7 of the 8
deaths (Table 30). The high percentage of male deaths attributable to hunting
was unexpected given that harvest of males was extremely limited on the study
area. Four of 7 males were harvested off the study area, indicating that males
did not utilize the mine as a refuge as has been suggested for other deer
populations (Zagata and Haugen 1973, Kammermeyer and Marchinton 1976).
Rather, males were harvested by hunters as they increased mobility and left the
study area during rut.
Table 29. Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult male mule deer, Rosebud Mine / Colstrip, Montana,
June 1992-May 1995.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk
15
16
18
18
18
18
10
9
18
18
18
17
#
#
deaths censored
#
added
Survival
0
0
0
0
0
1
1
0
0
0
0
1
1
2
0
0
0
0
0
9
0
0
0
0
1.000
1.000
1.000
1.000
1.000
0.611
0.611
0.611
0.611
0.611
0.577
0.577
0
0
0
0
0
7
0
0
0
0
1
0
Confidence interval
1.000<S<
1:000 < S <
1.000<S<
1.000<S<
1.000<S<
0.435 < S <
0.375 < S <
0.362 < S <
0.435 < S <
0.435 < S <
0.404 < S <
0.399 < S <
1.000
1.000
1.000
1.000
1.000
0.787
0.847
0.860
0.787
0.787
0.751
0.756
99
Table 30. Seasonal causes of death for radiocollared yearling / adult male mule
deer, Rosebud Mine / Colstrip, Montana, June 1992-May 1995.
Deer ID
C028-92
C041-93
C046-93
C054-93
C056-93
C064-93
C093-93
C040-93
Age at death
5.5
4.5
5.5
4.5
6.5
4.5
1.5
5.5
years
years
years
years
years
years
years
years
Date of death
04
15
20
15
22
26
13
01
NOV
NOV
NOV
NOV
NOV
NOV
NOV
APR
Season3
Cause of death
Fall
Fall
Fall
Fall
Fall
Fall
Fall
Spring
Hunter harvest
Hunter harvest
Hunter harvest
Hunter harvest
Hunter harvest
Hunter harvest
Hunter harvest
Unknown
1993
1994
1993
1993
1993
1993
1994
1993
a Seasonal designations: summer, 01 June-31 August; fall, 01 September-30
November; winter, 01 December-28 February; and spring, 01 March-31 May.
Overall patterns of annual survivorship differed greatly between
radiocollared females and males (X2=11.115, DF=1, P<0.001) (Figure 12).
Female mortality occurred primarily during March and April, while male mortality
occurred almost exclusively during the November general gun hunting season.
I
.§
e
3
CO
CC
JNE
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
Month
Female
Male
Figure 12. Monthly rates of survival for adult female and male mule deer,
Rosebud Mine / Colstrip, Montana, June 1992-May 1995.
MAY
100
MANAGEM ENT IMPLICATIONS
Management In Relation to Habitat
The Federal Surface Mining Control and Reclamation Act of 1977 requires
that surface-mined land be reclaimed to a condition equal to or higher than its
pre-mined condition. The Montana Strip and Underground Mining Act of 1979
further requires that reclamation be established with a permanent, diverse
vegetative cover consisting principally of native plants capable of, among other
things, sustaining grazing pressure by wild and domestic ungulates. Accordingly,
most surface mines in Montana and the western United States consider
establishment and maintenance of viable wildlife populations, including mule
deer, a primary reclamation goal (Scott and Zimmerman 1984).
Consistent with past research (Clark and Medcraft 1986, Medcraft and
Clark 1986), mule deer on the Rosebud Mine / Colstrip study area used disturbed
sites, especially reclamation, extensively. Phillips et al. (1985) documented not
only use of mining areas by mule deer but population increases as mining
progressed. Growth of the population was attributed to the diversification of
topographic and vegetative habitat features resulting from surface-mining and
reclamation actions.
Wildlife biologists have long recognized the importance of habitat diversity
and interspersion of vegetation-cover types to wildlife distribution and abundance .
(Leopold 1933, King 1938, Trippensee 1948, Allen 1954). Habitat diversity is
101
important for mule deer. As concentrate selectors (Hofmann 1985) that evolved
'v
in forest-edge habitats (Putman 1988, Geist 1994b), deer are adapted to
consume a wide variety of forages that vary in nutritional quality both among
species and seasons (Cook 1972, Freeland and Janzen 1974). Accordingly,
mule deer tend to forage either among vegetation-cover types that contain a high
diversity of plant species (Doucet et al. 1995) or among a diverse mosaic of types
providing highly interspersed forage resources. Similar to forage, cover needs of
mule deer and the ability of cover to satisfy those needs vary seasonally, making
diversity of vegetative and physiographic types within home ranges a necessity
(Mackie 1994a). Thus, mule deer require a diversity of vegetation-cover types for
survival and reproduction, and preservation of diversity should be a principal goal
where management seeks to maintain or increase mule deer numbers (Pac et al.
1991).
Surface mining on the Rosebud Mine likely has enhanced both
topographic and vegetative diversity of the area. Establishment of large spoil
ridges (12.1-42.4 m in height with a 45-54 degree slope) and sub- / top-soil
stockpiles (3.6-13.6 m in height with a 45-54 degree slope) in areas with flat to
rolling topography has increased local relief substantially. Diversity and
interspersion of vegetation-cover types also have increased as reclamation fields
seeded with a wide variety of native and introduced forbs or alfalfa have been
established adjacent to native vegetation providing both cover and alternative
forage resources. Mule deer apparently have responded favorably to these
!
habitat modifications based on (1) population growth since 1974, (2) mule deer
102
use of and distribution among mining disturbance and reclamation vegetationcover types, (3) measures of mule deer physical condition and female
productivity, (4) measures of adult female and fawn survival, and (5) overall
population age / sex structure.
Diversity of native vegetation-cover types on the Rosebud Mine should be
maintained in areas where deer presence is desired. Use of reclamation by deer
likely would be much reduced if interspersed native vegetation-cover types were
eliminated through future mining actions. During this study, native riparian and
pine savannah types were used preferentially within home ranges of females,
while reclamation often was preferred only at the landscape level. These findings
suggest that reclamation was important in determining placement of home
ranges, but maintenance of home ranges (i.e., deer survival and reproduction)
was influenced primarily, by the presence of native vegetation-cover types.
Further, findings from this study indicated that native vegetation, principally pine
savannah, had high value as accessory habitat for deer with home ranges
centered on reclamation and other vegetation-cover types. Loss of accessory
areas situated in native vegetation could render useless home ranges established
in marginal habitats (Pac et al. 1991), reducing deer abundance in these areas.
A second management consideration involves the landscape-scale
creation and placement of mining disturbance and reclamation vegetation-cover
types. Clearly, economic considerations are of over-riding importance when
developing coal-extraction plans. However, results of this and other studies
(Phillips et al. 1985) indicate that conflicts between mule deer and humans should
103
be expected when mining, followed by reclamation, occurs adjacent to centers of
human population such as Colstrip. To avoid future deer-human conflicts, mining
plans developed prior to disturbance should be drafted with a final goal of low
deer abundance on reclaimed lands adjacent to urban / developed areas. This
goal could be attained through (1) reduction of vegetative diversity, particularly
forb diversity, on reclamation, (2) reduction of structural diversity of reclamation
vegetation via elimination of shrubs, and (3) elimination of native vegetation,
especially preferred pine savannah and riparian vegetation-cover types, in areas
adjacent to reclamation. These actions would reduce the availability of preferred
diet items, required cover, and overall landscape heterogeneity, resulting in
habitat less conducive to survival and reproduction of mule deer within or
adjacent to human development.
Management In Relation to Humans
Given mule deer population responses to surface mining, the proximity of
Colstrip to the Rosebud Mine, and the observed use of Colstrip by. mule deer
inhabiting the Rosebud Mine, management also should consider immediate
means of minimizing current negative interactions between mule deer and the
residents of Colstrip. Observed negative deer-human interactions in Colstrip
during this study included deer-vehicle collisions, deer damage to gardens and
ornamental plantings, and direct physical contact between deer and humans I
pets.
Techniques used elsewhere to reduce the probability of such occurrences
104
have included the use of (1) guard dogs to frighten deer (Beringer et al. 1994), (2)
surgical sterilization or administration of contraceptive drugs to reduce deer
productivity (Turner et al. 1992, Frank et al. 1993), (3) preferred plantings to shift
deer distribution away from negatively impacted areas (Baron et al. 1966,
Campbell 1974, Nielsen et al. 1982), (4) frightening devices to scare deer away
from target areas (Dolbeer et al. 1994), (5) roadside improvements to warn
drivers of deer presence or frighten deer away from roadsides (Reed et al. 1982,
Schafer and Penland 1985), (6) chemical repellents to make plantings
unpalatable to deer (Conover 1984, Conover and Kania 1987), (7) physical
barriers to make property inaccessible to deer (Caslick and Decker 1979, Porter
1983, Palmer et al. 1985, Hygnstrom and Craven 1988), (8) live capture and
translocation to remove problem deer (Palmer et al. 1980, Ishmael and Rongstad
1984, Brush and Ehrenfeld 1991), and (9) selective harvest of deer or deer
groups to directly reduce deer abundance (Porter et al. 1991, Deblinger et al.
1992). These techniques have met with varying levels of success, but most are
deemed impractical due to high cost, unreliability, unacceptable levels of risk to
human safety, the tendency to simply shift the problem or create new problems
elsewhere, and the need to continually repeat the management action (O ’Bryan
and McCullough 1985, Crockett and Green 1986, Bryant and Ishmael 1991,
Wright 1993).
Methods that have consistently reduced deer damage to human property
or threats to human safety include (1) elimination I reduction of deer via intensive
removal programs and (2) exclusion of deer from areas of concern via
105
construction of physical barriers (Craven 1987). Removal via intensive harvest of
deer in Colstrip is impractical because of concerns for human safety. Removal
via live capture / translocation of deer is impractical due to high cost, poor
survivorship of translocated deer, and failure to address the initial need for
removal. Construction or improvement of physical barriers to impede movement
of deer into Colstrip, however, would require a one-time cost only and address
the goal of reducing deer abundance in Colstrip over time.
General observations and radiotelemetry data from this study indicated
deer seen in Colstrip made movements, principally crepuscular and nocturnal,
into the city from either the Rosebud Mine to the west or Montana Power
Company lands to the east. Most deer did not, however, establish home ranges
exclusively or even predominantly in Colstrip. Accordingly, the potential to
impede movements of deer into Colstrip exists. Deer-proof fences may
dramatically reduce deer abundance in Sections A, B, and C if established along
(1) the northern boundary of section B, (2) the western boundaries of sections A
and B, (3) the southern boundary of section A, and (4) the eastern boundaries of
sections A and C. Although fences currently are in place along the southern
boundary of section A and the eastern boundaries of sections A and C, they are
not maintained to the extent necessary to prevent deer from moving into and out
of Colstrip. Establishment of deer-proof fences in these areas would not only
reduce deer abundance and associated depredation problems in the southern
sections of Colstrip (sections A, B, and C), but likely also would reduce deervehicle collisions along State Highway 39 where it passes through Colstrip.
106
Because deer observed in urban areas were primarily those with home
ranges centered on the Rosebud Mine peripheral to Colstrip, selective deer
harvest on mining and reclamation areas immediately surrounding Colstrip could
further reduce deer presence in the city. The greatest known single-day distance
traveled by a deer to reach Colstrip from a point on the Rosebud Mine was
slightly over 1.7 km, although most deer traveled less than 800 m. Thus, harvest
of deer through regulated hunting or direct control, focused within a 2-km zone
surrounding Colstrip might also reduce the abundance of deer that have
established patterns of urban use. Because successful habitat-use strategies are
passed from generation to generation among members of matrilineal groups
(Dasmann and Taber 1956, Gruell and Papez 1963, Zalunardo 1965, Pac et al.
1991), elimination of deer, particularly females, with a history of urban use should
reduce “population knowledge” of Colstrip as a seasonably attractive habitat and
decrease the frequency of deer-human conflicts in the future.
Strategies designed to reduce deer abundance in and around Colstrip may
not be successful, however, if dispersing deer continually are attracted to
vegetation within Colstrip. To reduce the probability that deer will even attempt to
reestablish patterns of urban use within Colstrip, residents should be encouraged
to ( I ) landscape yards with plants unpalatable to deer, and (2) construct deerproof fencing around individual ornamental plants and / or garden plots.
Future Research Needs
This study broadly examined mule deer population characteristics and use
107
I
of habitats at a time when generally favorable environmental conditions led to
high mule deer numbers on the Rosebud Mine / Colstrip study area as well as
other portions of eastern Montana. Further, as of completion of this study,
W E C O had obtained bond release on none of its reclamation. Thus, these lands
were not subject to uses (e.g., livestock grazing and general hunter harvest of
deer) they will receive upon bond release. As mining is completed and
reclamation follows, topographic diversity provided by surface mining will
decrease due to removal of spoil and soil stockpiles and open mining pits.
Vegetative diversity derived from mining disturbance and reclamation also should
decrease as vegetative characteristics of reclaimed lands become increasingly
similar to surrounding native grassland and mixed shrub vegetation types.
Accordingly, the complete, long-term impacts of surface mining and reclamation
procedures on mule deer populations may only be assessed by monitoring deer
population characteristics and habitat use beyond the time of bond release when
reclaimed lands return to private ownership and management.
108
REFERENCES CITED
109
Aasheim1 R. 1980. Colstrip: a pleasant surprise. Montana Outdoors 11:16-17.
Ackerman, B. B., F. A. Leban1 M. D. Samuel, and E. 0 . Garton. 1989. User’s
manual for program home range. Univ. Idaho, For., Wildl., and Range
Exp. Stn., Tech. Rep. 15, Moscow. 79pp.
A lle n .D .L . 1954. Our wildlife legacy. Funk & Wagnalls, New York, N.Y.
422pp.
Anderson, A. E., and D. C. Bowden. 1972. Indices of carcass fat in a Colorado
mule deer herd. J. Wildl. Manage. 36:579-594. .
Arnold, G. W . 1985. Regulation of forage intake. Pages 81-101 in R. J.
Hudson, and R. G. White, eds. Bioenergetics of wild herbivores. CRC
Press, Inc., Boca Raton, Fla.
Bailey, N. T. J. 1951. On estimating the size of mobile populations from
recapture data. Biometrika 38:293-306.
Ball, S. K. 1987. Mule deer use of agricultural lands adjacent to Missouri River
Breaks habitat. Page 57 in A. Christensen, ed. Proc. Montana Chap.,
The Wildl. Soc., Kalispell.
Baron, F. J., E. C. Nord, A. B. Evanko, and W . J. Makel. 1966. Seeding conifers
and buffer crops to reduce deer depredation. Pacific Southwest For.
Range Exp. Stn., U S. For. Serv. Res. Note PSW^I 00. 8pp.
Barrett, M. W ., J. W . Nolen, and L. D. Roy. 1982. Evaluation of a hand-held netgun to capture large mammals. Wildl. Soc. Bull. 10:108-114.
Batchelor, R., M. Erwin, R. Martinka, D. McIntosh, R. Pfister, E. Schneegas, J.
Taylor, and K. Walther. 1982. A taxonomic classification system for
Montana riparian vegetation types: an interagency approach to classifying
Montana’s riparian ecosystems. Montana State Rural Areas Dev. Comm.,
Wildl., Subcomm,, Riparian Program Team, Bozeman, Mont. 13pp.
Beasom, S. L., W. Evans, and L. Temple. 1980. The drive net for capturing
western big game. J. Wildl. Manage. 44:478-480.
Beier1P. 1987. Sex differences in quality of white-tailed deer diets. J. Mammal.
68:323-329.
----- , and D. R. McCullough. 1988. Motion-sensitive radio collars for estimating
white-tailed deer activity. J. Wildl. Manage. 52:11-13.
110
Bellantoni, E. S., P. R. Krausman1 and W . W. Shaw. 1993. Desert mule deer
use of an urban environment. Trans. North Am. Wildl. And Nat. Resour.
C onf 58:92-101.
Beringer1 J., L. P. Hansen, R. A. Heinen1 and N. F. Giessman. 1994. Use of
dogs to reduce damage by deer to a white pine plantation. Wildl. Soc.
Bull. 22:627-632.
Blaxter1 K. L 1 R. B. Kay1 G. M. Sharman1 J. M. Cunningham, and W. J. Hamilton.
1974. Farming the red deer. H.M. Stationary Off., London, U.K. 93pp.
B onham .C .D . 1989. Measurements for terrestrial vegetation. John Wiley &
Sons, New York, N Y. 338pp.
Booth, W . E. 1950. Flora of Montana Part I: conifers and monocots. Montana
State Coll., Bozeman. 232pp.
----- , and J. C. Wright. 1962. Flora of Montana Part II: Dicotyledons. Montana
State Coll., Bozeman. 280pp.
Boukhout1 L. W . 1972. The behavior of mule deer (Odocoileus hemionus
hemionus) in winter in relation to social and physical environment. M S.
Thesis, Univ. Calgary. 124pp.
Bowyer1 R. T. 1984. Sexual segregation in southern mule deer. J. Mammal.
65:410-417.
— . 1986. Habitat selection by southern mule deer. Calif. Fish and Game
72:153-169.
— , and V. C. Bleich. 1984. Effects of cattle grazing on selected habitats of
southern mule deer. Calif. Fish and Game 70:240-247.
Brown, R. D. 1994. The nutrients for survival and growth. Pages 203-207 in D.
Gerlach et al./eds. Deer. Stackpole Books, Mechanicsburg1 Pa.
Brush, C. C., and D. W. Ehrenfeld. 1991. Control of white-tailed deer in nonhunted reserves and urban fringe areas. Pages 59-66 in L. W. Adams.and
D. L. Leedy1 eds. Wildlife conservation in metropolitan environments.
Natl. Inst. Urban Wildl., Columbia, Md.
Bryant, B. K., and W . Ishmdel. 1991. Movement and mortality patterns of
resident and translocated suburban white-tailed deer. Pages 53-58 in Li
W . Adams and D. L. Leedy1 eds. Wildlife conservation in metropolitan
environments. Natl. Inst. Urban Wildl., Columbia, Md.
111
Byers, C. R:, R. K. Steinhorst, and P. R. Krausman. 1984. Clarification of a
technique for analysis of utilization-availability data. J. Wildl. Manage.
48:1050-1053.
Campbell, D. L. 1974. Establishing preferred browse to reduce damage to
Douglas-fir seedlings by deer and elk. Pages 187-192 in H. C. Black, ed.
Wildlife and forest management in the Pacific Northwest. Oregon State
Univ., Corvallis.
Caslick, J. W ., and D. J. Decker. 1979. Economic feasibility of a deer-proof
fence for apple orchards. Wildl. Soc. Bull. 7:173-175.
Chalmers, B. A., and M. W . Barrett. 1982. Capture myopathy. Pages 84-94 in
G. L. Hoff and J. W . Davis, eds. Noninfectious diseases of wildlife. Iowa
State Univ. Press, Ames.
Chamrad, A. D., and T. W . Box. 1964. A point frame for sampling rumen
contents. J. Wildl. Manage. 28:473-477.
Cheatum, E. I . 1949. The use of corpora Iutea for determining ovulation
incidence and variations in fertility of the white-tailed deer. Cornell Vet.
39:282-291.
Clark, W . R., and J. R. Medcraft. 1986. Wildlife use of shrubs on reclaimed
surface-mined land in northeastern Wyoming. J. Wildl. Manage. 50:714718.
Clutton-Brock, T. H., F. E. Guinness, and S. D. Albon. 1982. Red deer: behavior
and ecology of two sexes. Univ. Chicago Press, Chicago, III. 378pp.
Cochran, W . W . 1980. Wildlife telemetry. Pages 507-520 in S. D. Schemnitz,
ed. Wildlife management techniques manual. The Wildlife Society,
Washington, D C.
Conner, M. C., E. C. Soutiere, and R. A. Lancia. 1987. Drop-netting deer: costs
and incidence of capture myopathy. Wildl. Soc. Bull. 15:434-438.
I
ConnoIIy1G -E . 1981. Assessing populations. Pages 287-345 jn 0 . C. Wallmo,,
ed. Mule and black-tailed deer of North America. Univ. Nebraska Press,
Lincoln.
Conover, M. R. 1984. Effectiveness of repellents in reducing deer damage in
nurseries. Wildl. Soc. Bull. 12:399-404.
112
— , and G. S. Kania. 1987. Effectiveness of human hair, BGR, and a mixture of
blood meal and peppercorns in reducing deer damage to young apple
trees. Proc. East. Wildl. Damage Control Conf 3:97-101.
Cook, C. W . 1972. Comparative nutritive values of forbs, grasses and shrubs.
Pages 303-310 In C. M. McKeII et al., tech. eds. Wildland shrubs: their
biology and utilization. U S. For. Serv., Intermountain For. and Range
Exp. Stn., Gen. Tech. Rep. INT-1, Ogden, U t
Cox, D. R., and D. Oakes. 1984. Analysis of survival data. Chapman and Hall,
New York, N Y. 201pp.
Craven, S. R. 1987. Deer. Pages D-23-33 in R. M. Timm, ed. Prevention and
control of wildlife damage. Nebraska Coop, Extension Serv., Univ. of
Nebraska, Lincoln.
Crockett, A. B., and R. A. Green. 1986. Mule deer in the city of Boulder: an
urban wildlife conflict and mangement alternatives. Pages 282-285 in R.
D. Comer et al, eds., Proc. II: Issues and technology in the management of
impacted wildlife. Thorne Ecol. Inst., Boulder, Colo.
Dasmann, R. F., and R. D. Taber. 1956. Behavior of Columbian black-tailed
deer with reference to population ecology. J. Mammal. 37:143-164.
Davis, D. E., and R. L. Winstead. 1980. Estimating the numbers of wildlife
populations. Pages 221-245 in S. D. Schemnitz, ed. Wildlife management
techniques manual. The Wildl. Soc., Washington, D.C.
Deblinger, R. D., J. J. Vaske, D. W. Rimmer, and M. P. Donnelly. 1992.
Ecological and social support for a controlled deer hunt. Pages 125-128 in
Proc. Northeast. Recreation Res. Symp., Northeast. For. Exp. Stn., U S.
For. Serv. Gen. Tech. Rep. NE-176.
Degayner, I. J., and P. A. Jordan. 1987. Skewed fetal sex ratios in white-tailed
deer: evidence of evolutionary speculations. Pages 178-188 in C. M.
Wemmer, ed. Biology and management of the Cervidae. Smithsonian
Inst. Press, Washington, D.C.
de Vos, J. C., Jr., C. R. Miller, and W. D. Ough. 1984. Habitat selection by mule
deer in a semi-urban area. Pages 128-131 in P. R. Krausman and N. S.
Smith, eds. Deer in the southwest: a symposium. School of Renewable
Nat. Resour., Univ. Arizona, Tucson.
113
Dolbeer1 R. A , N. R. Holler, and D. W . Hawthorne. 1994. Identification and
control of wildlife damage. Pages 474-506 in T. A. Bookhout1 ed.
Research and mangement techniques for wildlife and habitats. The Wildl.
Soc., Bethesda1 Md.
Dollhopf1 D. J., W . D. Hall, W . M. Schafer, E. J. DePuit1and R. L. Hodder.
1977a. Selective placement of coal strip mine overburden in Montana. I.
Data base. Montana Agric. Exp. Stn., Res. Rep. 128. Montana State
Univ., Bozeman. 109pp.
----- , ----- , C. A. Cull, and R. L. Hodder. 1977b. Selective placement of coal strip
mine overburden in Montana. II. Initial field demonstration. Montana
Agric. Exp. Stn., Res. Rep. 125. Montana State Univ., Bozeman. 98pp.
Dood1A. R. 1978. Summer movements, habitat use, and mortality of mule deer
fawns in the Missouri River Breaks, Montana. M.S. Thesis, Montana State
Univ., Bozeman. 55pp.
Doucet, C. M., E. D. Reid, K. L. Risen hoover, and C. A. Taylor. 1995. Foraging
response of ruminant herbivores to a gradient of resource availability.
Page 38 in The Wildlife Society second annual conference abstracts.
Portland, Oreg.
Downing, R. L, and B. S. McGinnes. 1969. Capturing and marking white-tailed
deer fawns. J. Wildl. Manage. 33:711-714.
Dusek, G. L. 1975. Range relations of mule deer and cattle in prairie habitat. J.
Wildl. Manage. 39:605-616.
----- , R. J. Mackie, J. D. Herriges, Fr., and B. B. Compton. 1989. Population
ecology of white-tailed deer along the lower Yellowstone River. Wildl.
Monogr. 104. 68pp.
Eberhardt, L. E., E. E. Hanson, and L. L. Cadwell. 1984. Movement and activity
patterns of mule deer in the sagebrush-steppe region. J. Mammal.
65:404-409.
Egan, J. L. 1957. Some relationships between mule deer and alfalfa production
in Powder River County, Montana. M.S. Thesis, Montana State Univ.,
Bozeman. 34pp.
Eustace, C. D. 1971. Mule deer food habits and browse use study. Montana
Dep. of Fish, Wildl., and Parks, Job Final Rep., Proj. W -130-R-1 and 2,
Job Number 1-7.1. 25pp.
114
Frank, E. S., S. L. Sajdak, J. A. Teare. 1993. Controlling urban white-tailed deer
by surgical sterilization. Page 245 in The Fifty-fifth Midwest Fish & Wildlife
Conference abstracts. St. Louis, Mo.
Freeland, W. J., and D. H. Janzen. 1974. Strategies in herbivory by mammals:
the role of plant secondary compounds. Am. Nat. 108:269-289.
Gavin, T. A., L. H. Suring, P. A. Vohs, and E. C. Meslow. 1984. Population
characteristics, spatial organization, and natural mortality in the Columbian
white-tailed deer. Wildl. Monogr. 91. 41pp.
Geist, V. 1990. Mule deer country. North Word Press, Inc., Minocqua, Wis.
176pp.
----- . 1994a. Social behavior. Pages 278-283 in D. Gerlach et al., eds. Deer.
Stackpole Books, Mechanicsburg, Pa.
-— . 1994b. Origin of the species. Pages 2-16 in D. Gerlach et al., eds. Deer.
Stackpole Books, Mechanicsburg, Pa.
Gillingham, M. P., and F. L. Bunnell. 1985. Reliability of motion-sensitive radio
collars for estimating activity of black-tailed deer. J. Wildl. Manage.
49:951-958.
Gilmer, D. S., V. B. Kuechle, and I. J. Ball, Jr. 1971. A device for monitoring
radio-marked animals. J. Wildl. Manage. 35:829-832.
----- , L. M. Cowardih, R. L. Duval, L. M. Mechlin, C. W. Schaiffer, and V. B.
Kuechle. 1981. Procdedures for the use of aircraft in wildlife biotelemetry
studies. U S. Fish & Wildl. Serv. Resour. Publ. 140. 19pp.
Glaze, R. E., and E. R. Keller. 1965. Geologic history of the Powder River
Basin. Am. Assoc. Petroleum Geol. Bull. 49:1893-1970.
Gruell, G. E., and N. J. Papez. 1963. Movements of mule deer in northeastern
Nevada. J. Wildl. Manage. 27:414-422.
Gysel, L. W ., and L. J. Lyon. 1980. Habitat analysis and evaluation. Pages 305327 in S. D. Schemnitz, ed. Wildlife management techniques manual.
The Wildlife Society, Washington, D C.
Hamlin, K. L. 1974. Ecological relationships of mule deer in the Bridger
Mountains, Montana, with special reference to daily and seasonal
movements. M.S. Thesis, Montana State Univ., Bozeman. 65pp.
115
— , and R. J. Mackie. 1989. Mule deer in the Missouri River Breaks, Montana:
population dynamics in a fluctuating environment. Montana Dep. Fish,
Wildl., and Parks, Fed. Aid in Wildl. Restor., Job Completion Rep., Proj.
W -120-R-7-18, Study Number BG-1.0. 401pp.
Hansen, P. L , G. R. Hoffman, A. J. Bjugstad. 1984. The vegetation of Theodore
Roosevelt National Park, North Dakota: a habitat type classification. U S.
For. Serv., Rocky Mountain For. and Range Exp. Stn., Gen. Tech. Rep.
RM-113. 35pp.
Happe, P. J. 1983. The use of suburban habitats by Columbian black-tailed
deer. M.S. Thesis, Oregon State Univ., Corvallis. 95pp.
Hawkins, R. E., L. DvMortogIio, and G. G. Montgomery. 1968. Cannon-netting
deer. J. Wildl. Manage. 32:191-195.
Heezen, K. L , and J. R. Tester. 1967. Evaluation of radiotracking by
triangulation with special reference to deer movements. J. Wildl. Manage.
31:124-141.
Hertzog, P. J. 1983. Response of native species to variable nitrogen,
phosphorus, and potassium fertilization on mine soils. M.S. Thesis,
Montana State Univ., Bozeman. 122pp.
Higgins, K. F., J. L. Oldemeyer, K. J. Jenkings, G. K. C.ambey, and R. F. Harlow.
1994. Vegetation sampling and measurement. Pages 567-591 in T. A.
Bookhout, ed. Research and mangement techniques for wildlife and
habitats. The Wildlife Society, Bethesda, Md.
Hofmann, R. R. 1985. Digestive physiology of the deer: their
morphophysiological specialisation and adaptation. Pages 393-407 in P.
F. Fennessy and K. R. Drew, eds. Biology of deer production. Royal Soc.
New Zealand.
Huegel, C. N., R. B. Dahlgren, and H. L. Gladfeleter. 1985. Use of doe behavior
to capture white-tailed deer fawns. Wildl. Soc. Bull. 13:287-289.
Hygnstrom, S. E., and S. R. Craven. 1988. Electric fences and commercial
repellents for reducing deer damage in cornfields, Wildl. Soc. Bull.
16:291-296.
Ihsle, H. B. 1982. Population ecology of mule deer with !emphasis on potential
impacts of oil and gas development along the east slope of the Rocky ,
Mountains, northcentral Montana. M.S. thesis, Montana State University,
Bozeman. 85 pp.
116
lhsle Pac, H. B., W . F. Kasworm, I . R. Irby, and R. J. Mackie. 1988. Ecology of
the mule deer, fOdocoileus hemionusl along the east front of the Rocky
Mountains, Montana. Can. Field-Nat. 102:227-236.
Irwin, L. L., and J. M. Peek. 1979. Shrub production and biomass trends
following five logging treatments within the cedar-hemlock zone of northern
Idaho. For. Sci. 25:415-426.
Ishmael, W . E., and 0 . J. Rongstad. 1984. Economics of an urban deer-removal
program. Wildl. Soc. Bull. 12:394-398.
Jackson, S. D. 1990. Ecology of mule deer on a sagebrush-grassland habitat in
northeastern Montana. M.S. Thesis, Montana State Univ., Bozeman.
111pp.
Jacobson, H. A. 1994. Feeding behavior. Pages 192-198 in D. Gerlach et al.,
eds. Deer. Stackpole Books, Mechanicsburg, Pa.
,
Jensen, W . F. 1992. Mule deer habitat use in the North Dakota badlands.
Prairie Nat. 24:97-108.
Julander, 0 , and J. B. Low. 1976. A historical account and present status of the
mule deer in the West. Pages 3-19 in G. W. Workman and J. B. Low,
eds. Mule deer decline in the west: a symposium. Utah State Univ., Coll,
of Nat. Resour., Agric. Exp. Stn., Logan.
Kammermeyer, K. E., and R. L. Marchinton. 1976. The dynamic aspects of deer
populations utilizing a refuge. Proc. Annu. Conf. Southeast. Assoc. Game
and Fish Comm. 29:466-475.
Kamps, G. F. 1969. White-tail and mule deer relationships in the Snowy
Mountains of central Montana. M.S. Thesis, Montana State Univ.,
Bozeman. 59pp. '
Kaplan, E. L., and P. Meier. 1958. Nonparametric estimation from incomplete
observations. J. Am. Stat. Assoc. 53:457-481.
Kenward, R. E. 1987. Wildlife radio tagging. Academic Press, London, U K.
222pp.
Kie, J. G., and M. White. 1985. Population dynamics of white-ailed deer
fOdocoileus virainianusl on the W elder Wildlife Refuge, Texas. The
Southwest. Nat. 30:105-118.
117
— , C. J. Evans, E. R. Loft, and J. W. Menke. 1991. Foraging behavior by mule
deer: the influence of cattle grazing. J. Wild!'. Manage. 55:665-674.
King, R. T. 1938. The essentials of a wildlife range. J. For. 36:457-464.
King, M. M., and H. D. Smith. 1980. Differential habitat utilization by the sexes of
mule deer. Great Basin Nat. 40:273-281.
Kjos, C. G., and W . W . Cochran. 1970. Activity of migrant thrushes as
determined by radio-telemetry. Wilson Bull. 82:225-226.
Knowles, C. J. 1975. Range relationships of mule deer, elk, and cattle in a restrotation grazing system during summer and fall. M S. Thesis, Montana
State Univ. 111pp.
Knudson, K., and L. Peterman. 1980. Wildlife and energy development:
Montana’s future options. Montana Outdoors 11:31-37.
Komberec, T. J. 1976. Range relationships of mule deer, elk, and cattle on a
rest-rotation grazing system during winter and spring. M.S. Thesis,
Montana State Univ. 79pp.
Kraft, S. 1987. Ecology of mule deer in the upper Missouri River Breaks,
Montana. M.S. Thesis, Univ. Montana, Missoula. 83pp.
Kucera, T. E. 1978. Social behavior and breeding system of the desert mule
deer. J. Mammal. 59:463-476.
Kufeld, R. C., 0 . C. Wallmo, and C. Feddema. 1973. Foods of the Rocky
Mountain mule deer. U S. Dep. Agric., For. Serv. Resear. Paper RM-111,
Fort Collins, Colo. 31pp.
Leopold, A. 1933. Game management. Univ.. Wisconsin Press, Madison.
481pp.
Lewis, B. D., and R. S. Roberts. 1977. Geology and water-yielding
characteristics of rocks of the northern Powder River Basin, southeastern
Montana. U S. Geol. Surv. Open-file Rep. 77-75. 22pp.
Linsdale, J. M., and P. Q. Tomich. 1953. A herd of mule deer. Univ. California
Press, Los Angeles. 567pp.
Loft, E. R. 1988. Habitat and spatial relationships between mule deer and cattle
in a Sierra Nevada forest zone. Ph D. Thesis, Univ. California, Davis.
144pp.
118
— , J. W . Menke1 and J. G. Kie. 1991. Habitat shifts by mule deer: the
influence of cattle grazing. J. Wildl. Manage. 55:16-26.
----- , ----- , and R. C. Bertram. 1987. Influence of cattle stocking rate on the
structural profile of deer hiding cover. J. Wildl. Manage. 51:655-664.
Lovell, D. S. 1992. Nonseeded species invasion of twelve revegetated surface
mined sites at Colstrip, Montana. M S. Thesis, Montana State Univ.,
Bozeman. 123pp.
Mackie, R. J. 1970. Range ecology and relations of mule deer, elk, and cattle in
the Missouri River Breaks, Montana. Wildl. Monogr. 20. 79pp.
----- . 1976. Interspecific competition between mule deer, other game animals
and livestock. Pages 49-54 in G. W . Workman and J. B. Low, eds. Mule
deer decline in the west: a symposium. Utah State Univ., Coll, of Nat.
Resour., Agric. Exp. Stn., Logan.
----- . 1978. Impacts of livestock grazing on wild ungulates. Trans. North Am.
Wildl. and Nat. Resour. Conf. 43:462-476.
— . 1983. Natural regulation of mule deer populations. Pages 112-125 in D. S.
Eastman et al., eds. Natural regulation of wildlife populations. Univ. Idaho
For., Wildl., and Range Exp. Stn., Moscow.
----- . 1994a. Mule deer habitat. Pages 286-296 in D. Gerlach et al., eds. Deer.
Stackpole Books, Mechanicsburg, Pa.
----- . 1994b. Reacting to weather. Pages 297-300 in D. Gerlach et al., eds.
Deer. Stackpole Books, Mechanicsburg, Pa.
----- . 1994c. Populations and habitat. Pages 322-328 in D. Gerlach et al., eds.
Deer. Stackpole Books, Mechanicsburg, Pa.
----- . 1994d. Longevity. Pages 328-329 in D. Gerlach et al., eds. Deer.
Stackpole Books, Mechanicsburg, Pa.
----- , and D. F. Pac. 1980. Deer and subdivisions in the Bridger Mountains,
Montana. Proc. Annu. Conf. West. Assoc. Fish and Wildl. Agencies
60:517-526.
— , G. L. Dusek, K. L. Hamlin, H. E. Jorgensen, D. F. Pac, and A. K. Wood.
1985. Management recommendations. Pages 1-8 in Montana deer
studies. Job Prog. Rep., Fed. Aid Proj. W -120-R-16. Montana Dep. Of
Fish, Wildl., and Parks, Helena.
119
Main, M. B. 1994a. Sexual segregation among ungulates: research and review.
Ph D. Thesis, Oregon State Univ., Corvallis. 120pp.
----- . 1994b. Segregation of the sexes. Pages 269-277 in D. Gerlach et al., eds.
Deer. Stackpole Books, Mechanicsburg, Pa.
----- , and B. E. Coblentz. 1991. Sexual segregation among ungulates: a critique.
Wildl. Soc. Bull. 18:204-210.
Marchinton, R. L , and D. H. Mirth. 1984. Behavior. Pages 129-168 in L. K.
Halls, ed. White-tailed deer: ecology and management. Stackpole Books,
Harrisburg, Pa.
Martinka, C. J. 1978. Ungulate populations in relation to wilderness in Glacier
National Park, Montana. Trans. North Am. Wildl. and Nat. Resour. Conf
43:351-357.
Mautz, W . W ., P. J. Pekins, and J. A. Warren. 1985. Cold temperature effects
on metabolic rate of white-tailed, mule, and black-tailed deer in winter coat.
Pages 453-457 in P. F. Fenhessy and K. R. Drew, eds. The biology of
deer production. The Royal Soc. of New Zealand, Bull. 22.
McCullough, D. R. 1979. The George Reserve deer herd: population ecology of
a K-selected species. Univ. Michigan Press, Ann Arbor. 271pp.
McGregor, R. L 1986. Anacardiaceae, the gashew family. Pages 571-575 in T.
M. Barkley, ed. Flora of the Great Plains. Univ. Press of Kansas,
Lawrence.
M e c h ,L .D . 1983. Handbook of animal radio-tracking. Univ. Minnesota Press,
Minneapolis. 107pp.
Medcraft, J. R., and W . R. Clark. 1986. Big game habitat use and diets on a
surface mine in northeastern Wyoming. J. Wildl, Manage. 50:135-142.
Meyn, R. L., E. S. Sunberg, and R. P. Young. 1976. Research on reclamation of
surface mined lands at Colstrip, Montana: progress report 1975. Montana
Agric. Exp. Stn. Res. Rep. 101. Montana State Univ., Bozeman. 84pp.
Miller, F. L. 1970. Distribution patterns of black-tailed deer (Qdocoileus
hemionus columbianus) in relation to environments J. Mammal. 51:248260.
Mohr, C. O. 1947. Tables of equivalent populations of North American small
mammals. Am. Midi. Nat. 37:223-249.
120
Nams, V. O., and S. Boutin. 1991. W hat is wrong with error polygons? J. Wildl.
Manage. 55:172-176.
National Oceanic and Atmospheric Administration. 1961-1995. Climatological
data: annual summary, Montana. Vols. 64-98. Natl. Climatic Data Cent.,
Asheville, N.C.
Neu, C. W ., C. R. Byers, and J. M. Peek. 1974. A technique for analysis of
utilization-availability data. J. Wildl. Manage. 38:541-545.
Nielsen, D. G., M. J. Dunlap, and K. V. Miller. 1982. Pre-rut rubbing by white­
tailed bucks: nursery damage, social role, and management options.
Wildl. Soc. Bull. 10:341-348.
Nietfeld, M. T., M. W . Barrett, and N. Silvy. 1994. Wildlife marking techniques.
Pages 140-168 in T. A. Bookhout, ed. Research and management
techniques for wildlife and habitats. The Wildlife Society, Bethesda, Md.
Nyberg, H. E. 1980. Distribution, movements and habitat use of mule deer
associated with the Brackett Creek winter range, Bridger Mountains,
Montana. M S. Thesis, Montana State Univ., Bozeman. 106pp.
O ’Bryan, M. K., and D. R. McCullough. 1985. Survival of black-tailed deer
following relocation in California. J. Wildl. Manage. 49:115-119.
O ’Connor, K. S. 1987. Ecology of white-tailed deer and mule deer in agricultural
lands in the Gallatin Valley, Montana. M.S. Thesis, Montana State Univ.,
Bozeman. 67pp.
Costing, H. J. 1956. The study of plant communities: an introduction to plant
ecology. W . H. Freeman and Company, San Francisco, Calif. 440pp.
Ordway, L. L , and P. R. Krausman. 1986. Habitat use by desert mule deer. J.
Wildl. Mange. 50:677-683.
Ozoga, J. J., and L. J. Verme. 1970. Winter feeding patterns of penned white­
tailed deer. J. Wildl. Manage. 34:431-439.
— , and — . 1982. Reproductive and physical characteristics of a
supplementally-fed white-tailed deer herd. J. Wildl. Manage. 46:281-301.
— , — , and C. S. Bienz. 1982. Parturition behavior and territoriality in white­
tailed deer: impact on neonatal mortality. J. Wildl. Manage. 46:1-11.
121
Pac, D. F., R. J. Mackie, and H. E. Jorgensen. 1991. Mule deer population
organization, behavior and dynamics in a northern Rocky Mountain
environment. Montana Dep. of Fish, Wildl., and Parks, Fed. Aid in Wildl.
Restor., Job Completion Rep., Proj. W -120-R-7-18, Study Number BG-1.2.
316pp.
Packer, P. E. 1974. Rehabilitation potentials and limitations of surface-mined
land in the northern Great Plains. U S. For. Serv. Gen. Tech. Rep. INT14, Intermountain For. and Range Exp. Stn., Ogden, Ut. 43pp.
Palmer, D. T., D. A. Andrews, R. 0 . Winters, and J. W. Francis. 1980. Removal
techniques to control an enclosed deer herd. Wildl. Soc. Bull. 8:29-33.
Palmer, W. L , J. M. Payne, R. G. Wingard, and J. L. George. 1985. A practical
fence to reduce deer damage. Wildl. Soc. Bull. 13:240-245.
Parker, K. L. 1994. The cost of living. Pages 306-314 in D. Gerlach et al., eds.
Deer. Stackpole Books, Mechanicsburg, Pa.
Payne, G. F, 1973. Vegetative rangeland types in Montana. Montana Agric.
Exp. Stn. Bull. 671. Montana State Univ., Bozeman. 15pp.
Phillips, R. L, D. E. Biggins, and A. B. Hoag. 1985. Coal surface mining and
selected wildlife: a 10-year case study near Decker, Montana. Pages 235245 in R. D. Comer, et al., eds. Proc. II: Issues and technology in the
management of impacted wildlife. Thorne Ecol. Inst., Boulder, Colo.
Plantenberg, P. L. 1983. Factors affecting vegetation development on mined
land at Colstrip, Montana. M S. Thesis, Montana State Univ., Bozeman.
121pp.
Pollock, K. H., S. R. Winterstein, C. M. Bunck, and P. D. Curtis. 1989. Survival
analysis in telemetry studies: the staggered entry design. J. Wildl.
Manage. 53:7-15.
Pond, D. B., and B. W . O ’Gara. 1994. Chemical immobilization of large
mammals. Pages 125-139 in T. A. Bookhout, ed. Research and
management techniques for wildlife and habitats. The Wildlife Society,
Bethesda, Md.
Porter, W . F. 1983. A baited electric fence for controlling deer damage to
orchard seedlings. Wildl. Soc. Bull. 11:325-327.
— , and K. E. Church. 1987. Effects of environmental pattern on habitat
analysis. J. Wildl. Manage. 51:681-685.
122
—
N. E. Mathews, H B. Underwood, R. W . Sage, and D. F. Behrend. 1991.
Social organization in deer: implications for localized management.
Environ. Manage. 15:809-814.
----- _----- , ----- , ----- , a n d ----- . 1992. The gas molecule theory: dispersal and
management of white-tailed deer. Page 79 in R. D. Brown, ed. The
biology of deer. Springer-Verlag, New York, N Y.
Putman, R. 1988. The natural history of deer. Cornell Univ. Press, Ithaca, N Y.
191pp.
Reed, D. F. 1981. Conflicts with civilization. Pages 509-535 in 0 . C. Wallmo,
ed. Mule and black-tailed deer of North America. Univ. Nebraska Press,
Lincoln.
— , T. D. I. Beck, and T. N. Woodard. 1982. Methods of reducing deer-vehicle
accidents: benefit-cost analysis. WNdI. Soc. Bull. 10:349-354.
Rees, J. W ., R. A. Kainer, and R. W. Davis. 1966. Chronology of mineralization
and eruption of mandibular teeth in mule deer. J. Wildl. Manage. 30:629631.
Riley, S. J. 1982. Survival in d behavior of radio-collared mule deer fawns during
summers, 1978-1980, in the Missouri River Breaks, Montana. M.S.
Thesis, Montana State Univ., Bozeman. 59pp. — , and A. R. Dood. 1984. Summer movements, home range, habitat use, and
behavior of mule deer fawns. J. Wildl. Manage. 48:1302-1310.
Riney, T. 1955. Evaluating condition of free-ranging red deer (Cervus elaohusl
with special reference to New Zealand. New Zealand J. Sci. Technol.
36:429-463.
Robinette, W . L. 1966. Mule deer home range and dispersal in Utah. J. Wildl.
Manage. 30:335-349.
— , D. A. Jones, G. Rogers, and J. S. Gashwiler. 1957. Notes on tooth
development and wear for Rocky Mountain mule deer. J. Wildl. Manage.
21:134-153.
— , C. H. Baer, R, E. Fillmore, and C. E. Knittle. 1973. Effects of nutritional
change on captive mule deer. J. Wildl. Manage. 37:312-326.
123
Sadleir, R. M. F. S. 1987. Reproduction of female Cervids. Pages 123-144 in C.
M. Wemmer, ed. Biology and management of the Cervidae. Smithsonian
Inst. Press, Washington, D C.
Samuel, M. D., and M. R. Fuller. 1994. Wildlife radiotelemetry. Pages 370-418
in T. A. Bookhout, ed. Research and management techniques for wildlife
and habitats. The Wildlife Society, Bethesda, Md.
SAS Institute, Inc. 1987. SAS system for elementary statistical analysis.
Version 5, SAS Inst., Inc., Cary, N.C. 416pp.
Scarbrough, D. L , and P. R. Krausman. 1988. Sexual segregation by desert
mule deer. The Southwest. Nat. 33:157-165.
.
Schafer, J. A., and S. T. Penland. 1985. Effectiveness of swareflex reflectors in
reducing deer-vehicle accidents. J. Wild!. Manage. 49:774-776.
Schafer, W . M., G. A. Nielsen, D. J. Dollhopf, and K. Temple. 1979. Soil
genesis, hydrological properties, root characteristics, and microbial activity
of 1- to 50-year old strip mine spoils. U S. Dep. Agric., Environ. Prot.
Agency Interagency Energy / Environ. Res. and Dev. Rep. 600/7-79-100.
Industrial Environ. Res. Lab., Cincinnati, Oh. 212pp.
Schwarzkoph, W . F. 1973. Range use and relationships of mule deer on the
west slope of the Bridger Mountains, Montana. M.S. Thesis, Montna State
Univ., Bozeman. 65pp.
Scott, M. D., and G M. Zimmerman. 1984. Wildlife management at surface coal
mines in the Northwest. Wildl. Soc. Bull. 12:364-370.
Severson, K. E. 1981. Plains habitats. Pages 459-485 in O. C. Wallmo, ed.
Mule and black-tailed deer of North America. Univ. Nebraska Press,
Lincoln.
— , and A. V. Carter. 1978. Movement and habitat use by mule deer in the
northern Great Plains, South Dakota. Pages 466-468 in D. N. Hyder, ed.
Proc First International Rangelands Congress. Soc. Range Manage.,
Denver, Colo.
Short, H. L. 1981. Nutrition and metabolism. Pages 99-127 in 0 . C. Wallmo, ed.
Mule and black-tailed deer of North America. Univ. Nebraska Press,
Lincoln.
Smoliak1 S. 1956. Influence of climatic conditions on forage production of
shortgrass rangeland. J. Range Manage. 9:89-91.
124
Stansberry, B. J. 1991. Ecology of mule deer in coniferous forest habitat in
western / northwestern Montana. M S. Thesis, Montana State Univ.,
Bozeman. 64pp.
Sutherland, D. 1986. Poaceae, the grass family. Pages 1113-1235 in T. M .
Barkley, ed. Flora of the Great Plains. Univ. Press of Kansas, Lawrence.
Swenson, J. E., S. J. Knapp, and H. J. Wentland. 1983. Winter distribution and
habitat use by mule deer and white-tailed deer in southeastern Montana.
Prairie Nat. 15:97-112.
Taber, R. D., and R. F. Dasmann. 1954. A sex difference in mortality in young
Columbian black-tailed deer. J. Wildl. Manage. 18:309-315.
Trippensee, R. E. 1948. Wildlife management: upland game and general
principles. McGraw-Hill Book Co., Inc., New York, N.Y. 479pp.
Turner, J. W ., I. K. M. Liu, and J. F. Kirkpatrick. 1992. Remotely delivered
immunocontraception in white-tailed deer. J. Wildl. Manage. 56:154-157.
United States Bureau of the Census. 1992. Census of population and housing:
summary tape file 3A, 1990. (CD90-3A-15) (CD-ROM), Idaho, Montana,
Nevada, and Wyoming. Washington, D C.
— . 1994. Statistical abstract of the United States. 114th edition. U S. Dep. Of
Treasury, Dep. O f Commer., Bur. of Stat., Econ. And Stat. Adm.,
Government Printing Office, Washington, D C. 1011pp.
Verme, L. J. 1969. Reproductive patterns of white-tailed deer related to
nutritional planes. J. Wildl. Manage. 33:881-887.
----- . 1983. Sex ratio variation in Odocoileus: a critical review. J. Wildl. Manage.
47:573-582.
— , and J. J. Ozoga. 1981. Sex ratio of white-tailed deer and the estrus cycle.
J. Wildl. Manage. 45:710-715.
Veseth, R., and C. Montagne. 1980. Geologic parent materials of Montana soils.
MontanaAgric. Exp. Stn. Bull. 721. Montana State Univ., Bozeman.
117pp.
Vogel, W . 0 . 1983. The effects of housing developments and agriculture on the
ecology of white-tailed deer and mule deer in the Gallatin Valley, Montana.
M.S. Thesis, Montana State Univ., Bozeman. 86pp.
125
Wallmo, 0 . C. 1978. Mule and black-tailed deer. Pages 31-41 in J. L. Schmidt
and D. L. Gilbert, eds. Big game of North America: ecology and
management. Stackpole Books, Harrisburg, Pa.
Welch, J. G., and A. P. Hooper. 1988. Ingestion of feed and water. Pages 108116 in D. C. Church, ed. The ruminant animal: digestive physiology and
nutrition. Waveland Press, Inc., Prospect Heights, III.
White, M., F. F. Knowlton, and W. C. Glazener. 1972. Effects of dam-newborn
fawn behavior on capture and mortality. J. Wildl. Manage. 36:897-906. ,
White, G. C., and R. A. Garrott. 1990. Analysis of wildlife radio-tracking data.
Academic Press, Inc., San Diego, Calif. 383pp.
Wilkins, B. T. 1957. Range use, food habits, and agricultural relationships of
mule deer, Bridger Mountains, M ontana.. J. Wildl. Manage. 21:159-169.
Wood, A. K. 1986. Ecology of a prairie mule deer population. Ph D. Thesis,
Montana State Univ., Bozeman. 205pp.
— , R. J. Mackie, and K. L. Hamlin. 1989. Ecology of sympatric populations of
mule deer and white-tailed deer in a prairie environment. Montana Dep. of
Fish, Wildl., and Parks, Fed. Aid in Wildl. Restor., Proj. W -10O-R and W 120-R, Bozeman. 97pp.
Wright, R. G. 1993. Wildlife management inparks and suburbs: alternatives to
sport hunting. Renewable Resour. J. 11:18-22.
Zagata, M. D., and A. 0 . Haugen. 1973. Pilot Knob State Park: a winter deer
haven. Iowa State J. Res. 47:199-217.
Zalunardo, R. A. 1965. The seasonal distribution of a migratory mule deer herd.
J. Wildl. Manage. 29:345-351.
126
APPEN DIX
/
127
Table 31. Capture / status of mule deer, Rosebud Mine / Colstrip, Montana, March
1992-May 1995.
D e e rl.D .3
Capture date Sex
CO01-92*
C002-92
C003-92*
C004-92*
C005-92*
C006-92*
C007-92*
C008-92*
C009-92
C 0 10-92
C 0 11-92*
C 0 12-92*
C 0 13-92*
C 0 14-92*
C 0 15-92*
C 0 16-92*
C017-92
C 0 18-92
C 019-92
C020-92
C021-92
C022-92*
C023-92
C024-92*
C025-92
C026-92*
C027-92*
C028-92*
C029-92*
C030-92*
C031-92*
C032-92
C033-93*
C034-93*
C035-93*
C036-93*
C037-93*
C038-93*
C039-93*
C040-93*
06 MAR 92
06 MAR 92
06 MAR 92
06 MAR 92
07 MAR 92
07 MAR 92
07 MAR 92
07 MAR 92
07 MAR 92
07 MAR 92
07 MAR 92
07 MAR 92
07 MAR 92
07 MAR 92
07 MAR 92
08 MAR 92
19 JUN 92
11 JUN 92
1 9 JUN 92
22 JUN 92
2 2 JUN 92
30 JUN 92
30 JUN 92
30 JUN 92
01 JLY 92
01 JLY 92
01 JLY 92
13 JLY 92
08 DEC 92
08 DEC 92
08 DEC 92
31 DEC 92
11 JAN 93
1 1 JAN 93
1 1 JAN 93
11 JAN 93
1 1 JAN 93
1 1 JAN 93
11 JAN 93
11 JAN 93
F
M
F
F
F
F
F
F
M
M
F
F
F
F
F
F
M
M
M
M
F
F
M
F
M
F
M
M
F
F
F
M
F
F
F
F
F
F
F
M
Ageb
.
4.5
2.5
7.5
2.5
2.5
6.5
3.5
2.5
2.5
0.5
2.5
1.5
9.5
3.5
3.5
5.5
3.5
0.0
0.0
0.0
0.0
1.5
1.5
1.5
0.0
2.5
2.5
4.5
4.5
0.5
. 0.5
0.5
3.5
7.5
2.5
5.5
6.5
1.5
5.5
5.5
Status:31 May 1995
Censored: 19 N O V 94
Dead: study collection, 12 APR 93
Dead: malnutrition, 01 APR 94
Censored: 20 AUG 94
Censored: 15 DEC 94
Dead: myopathy, 09 MAR 92
Dead: coyote predation, 15 SPT 92
Censored: 15 DEC 94
Unknown
Unknown
Censored: 01 JLY 94
Dead: hunter harvest, 15 NOV 94
Censored: 16 JAN 94
Dead: vehicle collision, 15 MAR 93
Censored: 15 DEC 94
Censored: 01 MAY 94
Dead: hunter harvest, 14 NOV 92
Unknown
Unknown
Unknown
Unknown
t
Censored: 19 N O V 94
Unknown
Alive / transmitting
Unknown
Censored: 19 N O V 94
Censored: 19 N O V 94
Dead: hunter harvest, 04 NOV 93
Censored: 30 MAR 95
Dead: unknown, 15 JAN 93
Dead: unknown, 14 MAR 93
Unknown
Censored: 27 MAY 95
Dead: vehicle collision, 13 APR 95
Alive / transmitting
Alive / transmitting
Censored: 27 MAY 95
Dead: vehicle collision, 13 APR 93
Alive / transmitting
Dead: unknown, 01 APR 93
128
Table 31. Continued, 2 of 3.
Deer I D.
Capture date Sex
Agea
C041-93*
C042-93*
C043-93*
C044-93*
C045-93*
C046-93*
C047-93*
C048-93*
C049-93
C050-93*
C051-93*
C052-93*
C053-93*
C054-93*
C055-93*
C056-93*
C057-93*
C058-93*
C059-93*
C060-93*
C061-93*
C062-93*
C063-93*
C064-93*
C065-93*
C066-93*
C067-93
C068-93*
C069-93
C070-93*
(2071-93*
C072-93*
C073-93*
C074-93*
C075-93*
C076-93*
C077-93*
C078-93*
C079-93*
C080-93*
C081-93*
11 JAN
1 1 JAN
1 2 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
1 2 JAN
1 2 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
12 JAN
13 JAN
13 JAN
1 3 JAN
1 3 JAN
1 3 JAN
1 3 JAN
13 JAN
1 3 JAN
1 3 JAN
05 JUN
06 JUN
06 JUN
08 JUN
08 JUN
08 JUN
08 JUN
09 JUN
1 0 JUN
3.5
5.5
5.5
3.5
3.5
4.5
2.5
3.5
0.5
4.5
6.5
5.5
2.5
3.5
6.5
5.5
3.5
4.5
2.5
8.5
9.5
3.5
2.5
3.5
7.5
8.5
0.5
6.5
0.5
2.5
9.5
3.5
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
93
M
F
F
F
F
M
F
F
M
F
F
M
F
M
F
M
F
M
F
F
F
F
F
M
F
F
M
F
F
M
F
F
M
M
M
M
M
M
M
M
M
'
Status:31 May 1995
Dead: hunter harvest, 15 N O V 94
Censored: 15 DEC 94
Alive / transmitting
Alive / transmitting
Alive / transmitting .
Dead: hunter harvest, 20 NOV 93
Dead: unknown, 26 NOV 94
Alive / transmitting .
Unknown
Alive / transmitting
Alive / transmitting
Alive / transmitting
/ Alive / transmitting
Dead: hunter harvest, 15 NOV 93
Alive / transmitting
Dead: hunter harvest, 22 NOV 93
Alive / transmitting
Censored: 27 MAY 95
Alive / transmitting
Dead: coyote predation, 14 MAR 94
Alive / transmitting
Alive / transmitting
Censored: 15 N O V 94
Dead: hunter harvest, 26 NOV 93
Alive / transmitting
Dead: unknown, 01 MAR 93
Unknown
Censored: 30 JLY 93
Unknown
Censored: 15 DEC 94
Alive / transmitting
Dead: coyote predation, 15 DEC 94
Dead: coyote predation, 01 OCT 93
Dead: coyote predation, 07 JAN 94
Censored: 07 JAN 94
Dead: coyote predation, 13 JUN 93
Dead: coyote predation, 29 MAR 94
Censored: 19 N O V 93
Dead: coyote predataion, 21 JAN 94
Censored: 02 JLY 93
Censored: 21 SEP 93
129
Table 31. Continued, 3 of 3.
Deer I D.
Capture date Sex
C082-93*
C083-93*
C084-93
C085-93
C086-93*
C087-93*
C088-93*
C089-93*
C090-93*
C091-93*
C092-93*
C093-93*
C094-93*
C095-94*
C096-94*
C097-94*
C098-94*
C099-94*
C l 00-94*
C l 01-94*
C 102-94*
C 103-94*
C l 04-94*
C l 05-94*
C 106-94*
C 107-94*
C l 08-94*
1 0 JUN
1 0 JUN
1 0 JUN
1 3 JUN
1 3 JUN
14 JUN
14 JUN
14 JUN
14 JUN
1 4 JUN
1 4 JUN
1 4 JUN
1 4 JUN
10 JUN
10 JUN
10 JUN
10 JUN
11 JUN
11 JUN
11 JUN
11 JUN
11 JUN
1 2 JUN
1 2 JUN
1 3 JUN
14 JUN
17 JUN
93
93
93
93
93
93
93
93
93
93
93
93
93
94
94
94
94
94
94
94
94
94
94
94
94
94
94
F
F
F
M
F
F
M
F
M
M
M
M
F
M
M
F
F
M
. M
M
M
M
M
M
M
F
M
Status:31 May 1995
Agea
0.0
0.0
0.0
0.0
0.0
0.0
. 0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
'
Dead: coyote predation, 19 JUN 93
Censored: 15 DEC 94
Unknown
Dead: coyote predation, 30 JUN 93
Dead: coyote predation, 01 FEB 94
Censored: 15 NOV 94
Dead: unknown, 27 AUG 93
Censored: 06 JLY 93
Dead: coyote predation, 11 JLY 93
Censored: 24 DEC 93
Dead: vehicle collision, 23 JLY 93
Dead: hunter harvest, 13 N O V 94
Alive / transmitting
Dead: vehicle collision, 28 AUG 94
Dead: coyote predation, 18 FEB 95
Dead: coyote predation, 26 JAN 95
Alive / transmitting
Dead: coyote predation, 26 JAN 95
Dead: coyote predation, 22 JUN 94
Dead: coyote predation, 18 FEB 95
Alive / transmitting
Alive / transmitting
Censored: 27 MAY 95
Alive / transmitting
Alive / transmitting
Censored: 01 APR 95
Censored: 14 O C T 94
a Asterisk following identification number signifies the deer was radiocollared.
b Age assigned as age at capture (age of 0.0 signifies a neonate fawn).
130
Table 32. Monthly survival estimates and 95% confidence intervals for
radiocollared female and male mule deer fawns, Rosebud Mine / Colstrip1
Montana, June 1993-May 1994.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk
21
18
14
13
12
11
10
9
6
5
4
4
#
#
deaths censored
#
added
Survival
0
2
0
' 1
0
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0.857
0.762
0.707
0.707
0.649
0.649
0.649
0.504
0.420
0.336
0.336
0.336
3
2
1
0
1
0
0
2
1
1
0
0
Confidence interval
0.719 < S <
0.590 < S <
0.507 < S <
0.499 < S <
0.431 < S <
0.421 < S <
0.410 < S <
0.272 < S <
0.164<S<
0.096 < S <
0.068 < S <
0.068 < S <
0.996
0.934
0.908
0.915
0.866
0.876
0.887
0.736
0.676
0.576
0.605
0.605
Table 33. Monthly survival estimates and 95% confidence intervals for
radiocollared female and male mule deer fawns, Rosebud Mine / Colstrip,
Montana, June 1994-May 1995.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk
14
13
13
12
12
11
11
11
9
7
7
6
#
#
deaths censored
#
added
Survival
0
0
0
0
1
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0.929
0.929
0.857
0.857
0.857
0.857
0.857
0.701
0.545
0.545
0.545
0.545
1
0
1
0
0
0
0
2
2
0
0
0
Confidence interval
0.799 < S <
0.794 < S <
0.681 < S <
0.674<S<
0.674<S<
0.666 < S <
0.666 < S <
0.475 < S <
0.305 < S <
0.273<S<
0.273 < S <
0.251 < S <
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.928
0.786
0.818
0.818
0.840
131
Table 34. Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult female mule deer, Rosebud Mine / Colstrip,
Montana, June 1992-May 1993.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk
#
#
deaths censored
#
added
Survival
O
O
O
O
O
O
O
O
0.
0
0
0
2
1
0
0
0
0
1
28
0
0
0
0
1.000
1.000
1.000
0.933
0.933
0.933
0.933
0.933
0.933
0.890
0.868
0.868
12
14
15
15
14
14
14
15
43
43
41
40
O
O
O
1
O
O
O
O
O
2
1
O
Confidence interval
1.000 < S <
1.000<S<
1.000<S<
0.811 < S <
0.807 < S <
0.807 < S <
0.807<S<
0.811 < S <
0.861 < S <
0.802 < S <
0.772 < S <
0.771 < S <
1.000
1.000
1.000
1.000
1.000
1,000
1.000
1.000
1.000
0.978
0.965
0.966
Table 35. Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult female mule deer, Rosebud Mine / Colstrip,
Montana, June 1993-May 1994.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk
40
40
39
39
39
39
39
39
38
38
37
36
#
#
#
deaths censored added
Survival
0
0
0
0
0
0
0
0
0
0
0
0
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.974
0.947
0.947
,
0
0
0
0
0
0
0
0
0
1
1
0
0
1
0
0
0
0
0
1
0
0
0
1
Confidence interval
.
1.0 0 0 <
1 .0 0 0 <
1.000 <
1.0 0 0 <
1.0 0 0 <
t :o o o <
1.000 <
1.0 0 0 <
1.000 <
0.923 <
0 .8 7 7 <
0.876 <
S
S
S
S
S
s
S
S
S
S
S
S
<
<
<
<
<
<
<
<
<
<
<
<
1.000
1.000
1.000
1.000
1.000
lo c o
1.000
1.000
1.000
1.000
1.000
1.000
132
Table 36. Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult female mule deer, Rosebud Mine / Colstrip,
Montana, June 1994-May 1995.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk
35
38
37
36
36
36
29
23
23
23
22
21
#
#
deaths censored
#
added
Survival
O
1
1
O
O
5
5
O
O
1
O
2
3
O
O
O
O
O
O
O
O
O
O
O
1.000
1.000
1.000
1.000
1.000
0.944
0.912
0.912
0.912
0.912
0.870
0.870
O
O
O
O
O
2
1
O
O
O
1
O
Confidence interval
«
1.000
1.000
1,000
1.000
1.000
0.872
0.813
0.801
0.801
0.801
0.740
0.736
<S<
< S<
< S<
< S<
<S<
< S <
< S <
<S<
< S<
< S<
<S<
< S<
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
Table 37. Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult male mule deer, Rosebud Mine / Colstrip, Montana,
June 1992-May 1993.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk
0
0
2
2
2
2
2
2
11
11
11
10
#
#
#
deaths censored added
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
b
0
0
2
0
0
0
0
0
9
0
0
0
0
Survival
Confidence interval
NA
NA
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.909
0.909
NA
NA
1.000
1.000
1.000
1,000
1.000
1.000
1.000
1.000
1.000
1.000
1.000<S<
1.000 < S <
1.000 < S <
1.000 < S <
1.000<S<
1.000 < S <
1.000 < S <
1.000<S<
0.747 < S <
0.739 < S <
133
Table 38. Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult male mule deer, Rosebud Mine / Colstrip1 Montana,
June 1993-May 1994.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk
10
10
10
10
10
10
5
5
5
5
5
5
#
#
deaths censored
#
added
Survival
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.000
1.000
1.000
1.000
1.000
0.500
0.500
0.500
0.500
0.500
0.500
0.500
0
0
0
0
0
5
0
0
0
0
0
0
Confidence interval
1.000 < S <
1.000 < S <
1.000 < S <
1.000<S<
1.000<S<
0.281 < S <
0.190<S<
0.190 < S <
0.190 < S <
0.190 < S <
0.190<S<
0.190 < S <
1.000
1.000
1.000
1.000
1.000
0.719
0.810
0.810
0.810
0.810
0.810
0.810
Table 39. Monthly survival estimates and 95% confidence intervals for
radiocollared yearling / adult male mule deer, Rosebud Mine / Colstrip, Montana,
June 1994-May 1995.
Month
JUN
JLY
AUG
SPT
OCT
NOV
DEC
JAN
FEB
MAR
APR
MAY
#
at risk.
5
6
6
6
6
6
3
2
2
2
2
2
#
#
deaths censored
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
1'
1
0
0
0
0
1
#
added
Survival
1
0
0
0
0
0
0
0
0
0
0
0
1.000
1.000
1.000
1.000
1.000
0.667
0.667
0.667
0.667
0.667
0.667
0.667
Confidence interval
1.000<S<
1.000 < S <
1.000 < S <
1.000<S<
1.000<S<
0.359 < S <
0.231 < S <
0.133 < S <
0.133 < S <
0.133 < S <
0.133 < S <
0.133 < S <
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
134
—
I
Ridgetop
I W E C O R eclam ation (1 9 9 4 )
BS888SS Colstrip
+ D e e r group
IalsSii A rea not surveyed
Figure 13. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer and winter 1974.
_____________
Rldgetop
I
- I W E C O R eclam ation (1 9 94 )
BfififfiiSi Colstrlp
+ D e e r group
I i l s l i A rea not surveyed
Figure 14. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1975, and spring
1976.
135
+ D e e r group
I # : # # A re a not surveyed
Figure 15. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1976, and spring
1977.
+ D e e r group
i U A rea not surveyed
Figure 16. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1977, and spring
1978.
136
+
___________ i
"
Ridgetop
I.. - -J W E C O R e clam atio n (1 9 9 4 )
B888888 Colstrip
+ D e e r group
A rea not surveyed
Figure 17. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1978, and spring
1979.
—
I
Ridgetop
I W E C O R eclam ation (1 9 9 4 )
S8BS8 Colstrip
+ D e e r group
H A re a not surveyed
Figure 18. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1979, and spring
1980.
137
—
Rl dget op
I
I W E C O R e clam atio n (1 9 9 4 )
BSSSSSS Colstrlp
K-
+ D e e r group
I A rea not surveyed
Figure 19. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1980, and spring
1981.
++++
I
Rldgetop
I W E C O R e clam atio n (1 9 94 )
3888888 Colstrlp
+ D e e r group
A rea not surveyed
Figure 20. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1981, and spring
1982.
138
+ + +
Ridgetop
I
I W E C O R eclam ation (1 9 94 )
SSSSK Colstrip
+ D e e r group
lL:_3 A rea not surveyed
Figure 21. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1982, and spring
1983.
\
..... Ridgetop
I
I W E C O R e clam atio n (1 9 9 4 )
BSSSSft Colstrip
+ D e er group
I:::: :-:': N A rea not surveyed
Figure 22. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1983, and spring
1984.
139
' “ ' Ridgetop
I-------- 1W E C O R eclam ation (1 9 9 4 )
855888 Colsthp
__ + D e e r group
A rea not surveyed
Figure 23. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1984, and spring
1985.
_________________
Rldgetop
- 1W E C O R eclam ation (1 9 94 )
SBSSSSS Colstrlp
+ D e e r group
K m W A rea not surveyed
I
Figure 24. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1985, and spring
1986.
140
—
I
Ridgetop
-I W E C O R e clam atio n (1 9 9 4 )
Colstrlp
+ D e e r group
Figure 25. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1986, and spring
1987.
Ridgetop
I
I W E C O R eclam ation (1 9 9 4 )
8 8 8 ® Colstrip
+ D e e r group
Figure 26. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1987, and spring
1988.
141
O ^
+
++
Ridgetop
l W E C O R e clam atio n (1 9 94 )
3885889 colstrip
+ D e e r group
I
Figure 27. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1988, and spring
1989.
+ ++
" Ridgetop
I ■—J W E C O R eclam ation (19 94 )
8888558 colstrip
+ D e e r group
Figure 28. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1989, and spring
1990.
142
Kilometers
+ + +
Ridgetop
I
I W E C O R eclam ation (1 9 94 )
S888S8 colstrip
+ D e e r group
Figure 29. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1990, and spring
1991.
—
I
Rldgetop
I W E C O R eclam ation (19 94 )
SSSKKS Colstrip
+ D e e r group
Figure 30. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1991, and spring
1992.
143
Kilometers
+ + +
+ +
+
Ridgetop
I...... I W E C O R eclam ation (1 9 9 4 )
$85888 Colstrlp
+ D e e r group
Figure 31. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1992, and spring
1993.
Kilometers
+ + +
+ + +
+ + +
Ridgetop
+ +++ +
I
I W E C O R eclam ation (1 9 9 4 )
8888888 Colstrip
+ D e e r group
Figure 32. Distribution of mule deer groups on the Rosebud Mine permit area
based on fixed-wing aerial surveys, summer, fall, and winter 1993, and spring
1994.
MONTANA STATE UNIVERSITY LIBRARIES
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