HomeMy WebLinkAboutAPA1161Cover design: Wolverine tracks, Igloo Creek, Mount McKinley
National Park . Posterized from Kodachrome photo by F. Dean .
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REACTIONS OF WILDLIFE TO HUMAN ACTIVITY
ALONG t'10UNT ~1CKINLEY NATIONAL PARK ROAD
RECOiltr·1ENDED:
Chairman,Advisory Committee
~c,~
Acting Chairman,Proqram Wildlife
and'Fisheries
APPROVED:
Actin~Dean,Colleqe of Environmental Sciences
Date i
Vice-Chancellor for Research and Advanced Study
Date
ALASKA RESOURCES LIBRARY
U.s.DEPT.OF INTERIOR
REACTIONS OF \HLDLIFE TO HUMAN ACTIVITY
ALONG MOUNT MCKINLEY NATIONAL PARK ROAD
A
THESIS
Presented to the Faculty of the
University of Alaska in partial fulfillment
of the Requirements
for the Degree of
MASTER OF SCIENCE
By
J-,'
Diane Marie Tracy,B.S.
Fairbanks,Alaska
May 1977
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Abstract
To evaluate the impact of human activities on wildlife along
McKinley Park road,reactions of caribou,Moose,Dal1 sheep,brown
bears and red foxes were quantified during 100 trips along the road
and 300 hr of intensive ohservations in 1973-1974.Backcountry and
near-road mammal densities were determined in seven plots.Avoidance
was found only for some bears and foxes and,possibly,large bands of
caribou.Disturbances thwarted some road crossings ~y migrating sheep.
Disturbances decreased feeding and increased movements by caribou with-
..,
in 200 m of the road.For ungulates,females with young were most
easily disturbed;injuries to young may occur.Many individual animals
appeared habituated to human activities.All species exhibited 50%"no
visible response"between 50-100 m from the road.Few visible responses
were given by animals beyond 400 m.Loud noises or people out of
vehic1 es increased response strength for most speci es.r1any anima 1s
were attracted to the road.
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Acknowledgments
This study was funded by the National Park Service throuqh
Contracts CX-9000-4-0038 and CX-9000-5-0005.administered by the
Alaska Cooperative Park Studies Unit.I received a graduate stipend
and supplementary research funds through the National Science Foundation
Graduate Fellowship Program.
I would like to thank Dr.Frederick C.Dean for advice and en-
couragement during all phases of this study.Dr.Stephen F.Maclean,
Jr.and Dr.Peter C.lent gave much appreciated help during the plan-
ning stages of the study and provided critical reviews of the manuscript.
Their efforts are gratefully acknowledged.I \llou1d also like to thank
Dr.Samuel J.Harbo for assistance with the statistical analyses.
Special thanks go to Harold IIChi p ll Do"ming,Bowdy Train and
Patrick Va1kenburg,all of whom assisted me in the field.Without
their help this study could not have been conducted.Many National
Park Service personnel stationed at Hount r1cKinlev National Park aided
me while I was in the park.Gary Brown,Chief Ranger,and Steven Buskirk,
park biologist,were particularly cooperative and helpful,and I am very
grateful to them.
I am also very-indebted to Candy Anderson and Mary Slemmons for
putting in long hours of typing to produce the final manuscript.I
would 1ike to thank Audrey ~~agoun for he1 pi ng type the rough draft.
Finally,I thank my husband,Oouq1as Schamel,for endless patience,
encouragement and advice,for hours of typing and proofing,and for
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TABLE OF CONTENTS
INTRODUCTION
STUDY AREA
t~ETHODS . . •
RESULTS AND DISCUSSION
Visitation and Traffic Levels
Caribou (Rangifer tarandus)
Population History,Numbers,Composition,
Distribution • • • ..••.
Human Disturbance ..
General Avoidance
Reaction Distances .
Disturbance Frequencies
Relationshio of Reaction to Type of
Di sturbance'. . . • • . • . . . • .
Relationship of Reaction to Group Size .
Relationship of Reaction to Age and Sex
Relationship of Reaction to Behavior Before
Disturbance • . . . . • . .••.•..•
Relationship of Reaction to Time and
Season . . . . . . . . . . . . . . •
Relationship Of Reaction to Weather and
....Habitat '.
Effects of Disturbance on Activity Pattern
PAGE
1
4
8
16
16
22
22
30
30
34
40
41
47
49
51
53
56
57
Behavior Patterns Related to Disturbance.59
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Summary and General Discussion
vi
65
vii
Behavior Patterns Related to Disturbance.
Relationship of Reaction to Sex and Age
Relationship of Reaction to Season,Time,
Weather,Habitat • . • •
Behavior Before Disturbance
72
72
79
79
81
96
92
94
85
89
PAGE
...
..
. ... .. . .. .. . . . . .
Reaction Distances
General Avoidance
Relationship of Reaction to Type of
Disturbance 0 ••0 0 ••0 • 0
Population History,Numbers,Composition,
Di s tri buti on • • • •
Human Disturbance.
Moose (AZaes aZaes)
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General Avoidance
Reaction Distances •
Human Disturbance 0
105
109
109
111
100
100
105•eo • •.••
Population History,Numbers,Composition,
Distribution .0 0 0 0 0
Influence of Roadside Vegetation
Summary and General Discussion
Da11 Sheep (Ovis daZZi)
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Relationship of Reaction to Type of
"Disturbance 0 •••••••••
Relationship of Reaction to Sex and Age
Relationship of Reaction to S~ason,Weather,
Habitat and Experience •.0..0 0 0 0
Behavior Before Distu~bante
117
121
122
123
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Behavior Patterns Related to Disturbance •125
.. . .. . . ..... .. .
167
171
171
173
154
158
162
162
163
167
149
153
130
141
141
144
148
125
130
. .
... ..
. ... . . . ....
Reaction Distances ••
Reaction Distances
Disturbance Frequencies
General Avoidance
Behavior Patterns Related to Disturbance
Influence of Roadside Vegetation
Relationship of Reaction to Sex and Age
Relationship of Reaction to Behavior Before
Disturbance ••.•.••••••••.•
Relationship of Reaction to Type of
Di~turbance • • . • . • • • • .
Influence of Garbage ••••
"General Avoidance
Human Disturbance .•
Human Disturbance .
Population History,Numbers,Composition,
Distribution .•••
PopUlation History,Numbers,Composition,
Distribution
Summary and General Discussion
Summary and General Discussion
Brown Bear (Ursus arctos)
Red Fox (VuZpes vuZpes)
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Relationship of Reaction to Type of
Disturbance ••••••••••••... ..175
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Relationship of Reaction to Age and Sex 177
Use of Road as Travel Route 179
Behavior Patterns Related to Disturbance 180
Feeding of Foxes by Visitors.
Arctic Ground Squirrel (Spermophilus undulatus)
Collared Pika (Ochotona collaris).
182
183
184
184
189
189
190
193
194
195
195
..
..
.....
.. ..... .
... . . . . ..
Wolf (Canis lupus)
Lynx (Lynx canadensis)
Wolverine (Gulo gulo)•
Porcup i ne (Erethizon dorsatwn)
Snowshoe Hare (Lepus americanus)
Marmots (Manmota caligata)
Summary and General Discussion
Other r1amma 1s
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Short-eared Owl (Asio flammeus)•.
Golden Eagle (Aquila chrysaetos)
Long-tailed Jaegers (Stercorarius longicaudus)
Willow Ptarmigan (Lagopus lagopus)
. . ... ... . . . ..... ... ...
201
205
..",.I~O
196
196
197
198
198
201
...
...
. ......
Mew Gull (Larus canus)
Interacting Variables Affecting Responses
Birds
General Biological Concepts and the Responses of
Wildlife to Human Activities .••••'••
FINAL DISCUSSION
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Reaction Distances ..••
Type of Disturbance
Sex and Age . . .
Effects of Disturbance
Attraction of Wildlife to Road.
~1anagement Recommendati ons . . .
APPENDIX A.SUPPLEMENTARY DATA TABLES
LITERATURE CITED • • . • • • • • . . • .
•••
....
.....
PAGE
206
208
210
210
212
213
217
249
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LIST OF FIGURES
PAGE
Figure 4.Average daily number of vehicles passing east
plus west through the Savage River Guard Station
in 1974,~10unt ~1cKin1ey National Park,Alaska 19
Figure 5.Hourly traffic pattern during July 1974 along
the Igloo Canyon -Sable Pass section of the
road,Mount McKinley National Park,Alaska..20
Fig4re 1.~10unt ~kKin1ey National Park and features of the
road,1973-1974 . • • • • • • • •5
Figure 2.Location of study plots,1973-1974 • • • . • •9
Figure 3.Mount McKinley National Park annual visitation
levels,based on U.S.Department of Interior
(1974)report • • • • • • • • • • . • • • • •17
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U Figure 6.Reactions of caribou,observed at varying
distances from the road during shuttle-tour
trips,to the bus and visitors,1973-1974,
Mount McKinley National Park,Alaska ••••36
Figure 7.Reactions of caribou,excluding calves,to
traffic and visitors;comparison of shuttle-tour
observations and intensive observations,1973-
1974,t10unt McKinley National Park,Alaska.•••38
Figure 8.Reactions of caribou,observed within 200 m of
the road during shuttle-tour trips,to various
actions of the bus and.visitors,1973-1974,
Mount t1cKin1ey National Park,Alaska •.••
Figure 9.Reactions of moose,observed at varying
distances from the road durinq shuttle-tour
trips,to the bus and visitors,1973-1974,
'Mount ~1cKin1ey National Park,Alaska ••••
Figure 10.Reactions of moose,observed within 200 m of
th~'road during shuttle-tour trips,to various
actions of the bus and visitors,1973-1974,
Mount ~1cKin1ey National Park,Alaska ••
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44
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PAGE
Figure 11.Reactions of nall sheep,observed at varying
distances from the road during shuttle-tour
trips,to the bus and visitors,1973-1974,
Mount McKinley National Park,Alaska 113
Figure 12.Reactions of Dall sheep,observed within 200 m
of the road during shuttle-tour trips,to
various actions of the bus and visitors,1973-
1974;Mount McKinley National Park,Alaska.119
Figure 13.Reactions of brown bears,observed at varying
distances from the road during shuttle-tour
trips,to the bus and visitors,1973-1974,
Mount t1cKinley National Park,Alaska ••••••146
Figure 14.Reactions of brown hears,observed within 200 m
of the road durin~shuttle-tour trips,to
various actions of the bus and visitors,1973-
1974,~1ount McKinley National Park,Alaska ••151
xii
LIST OF TABLES
Table 4.Sex and age distributions of caribou observed
during shuttle-tour trips,~1ount r'1cKinley
National Park,Alaska ........•....~•
Table 3.Sizes of caribou groups observed during shuttle-
tour trips,excluding the migrating bands of
over 50 animals,Mount McKinley National Park,
Alas·ka ••••••••••••••••••••
Table 1.N~mbersof caribou observed during shuttle-tour
trips,excluding migrating bands of over 50
animals,Nount r'1cKinley National Park,Alaska.
Tabl~2.Sightings of migrating bands of Over 50 caribou
from the road,t10unt tkKinley National Park,
Alas'ka • • • • • • • • . • • •~• • • . • . • • •
28
26
31
29
23
PAGE
Average number of caribou observed in study
plots,Mount McKinley National Park,Alaska
Table 5.r
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Table 6.Reactions of caribou,observed at varying
distances from the road during shuttle-tour
trips,to the bus and visitors;percent of
animals and groups,1973-1974,~1ount r1cKinley
National Park,Alaska ••••••••••••35
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Table 7.
Table 8.
Table 9.
Reactions of caribou,at va~ying distances from
the road,to traffic and visitors,as observed
during shuttle-tour trips and during intensive
observations,calves excluderl,1973-1974,Mount
t~cKinley National Park,Alaska .•••.•••
Reactions of caribou,observed within 200 m of
the road during shuttle-tour trips,to various
actions of the bus and'visitors;percent of
animals and groups,1973-1974,Mount McKinley
National Park ••.••••••••••••••
Reactions of caribou,observed during shutt1e-
tour trips within 200 m of the road,in various
size groups to the bus and visitors,1973-1974,
Mount t1cKinley National Park,Alaska •••••
. .
37
43
48
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Table 10~Reactions of adult and calf caribou,observed
during intensive observations,to vehicles and
visitors,1974,t10unt t1cKinley National Park,
Alaska .
Table 11.Reactions of caribou,observed within 200 m of
the road during shuttle-tour trips and during
intensive observations,as related to behavior
before disturbance,1973-1974,Mount McKinley
National Park,Alaska ••••••••••
Table 12.Seasonal changes in reactions of caribou,
observed within 200 m of the road durinG
shuttle-tour trips,to the bus and visitors,
Mount McKinley National Park,Alaska •.•.•
Table 13.Behavior of caribou observed during shuttle-
tour trips and intensive observations,number
of animals exhibiting given lIinitialll behavior
when first sighted during shuttle-tour trips
and caribou-minutes of 9iven behavior during
intensive observations,1973-1974,Mount
tkKinley National Park,Alaska •.•••.•
Table 14.Maximu~behavioral reactions of caribou,
observed durina shuttle-tour and intensive
observations,to vehicles and visitors,1973-
1974,t·1ount tkKinley National Park,Alaska.
Table 15.Number of moose observed during shuttle-tour
trips,t'1ount tkKinley National Park,Alaska
Table 16.Sex and age distributions of moose observed
during shuttle-tour trips,Mount McKinley
National Park,Alaska •••..•.••••
Table 17.Sizes of moose groups observed during
shuttle-tour trips,Mount McKinley National
Park,Alaska ••..•••••.•••••.
Table 18.Average number of moose observed in study
plots,Mount McKinley National Park,Alaska
PAGE
50
52
55
58
60
73
75
78
80
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Table 19~
Table 20.
Table 21.
Table 22.
Table 23.
Table 24.
Table 25.
Table 26.
Table 27.
Reactions of moose,observed at varying
distances from the road during shuttle-tour
trips,to the bus and visitors;percent of
animals and groups,1973-1974,Mount McKinley
National Park,Alaska .••••••••••
Reactions of moose,observed within 200 m of
the road during shuttle-tour trips,to various
actions of the hus and visitors;percent of
animals and groups,1973-1974,r~ount r4cKin1ey
National Park,Alaska ••••••••••••
Reactions of various sex and age classes of
moose,observed during shuttle-tour trips,to
buses and visitors,1973-1974,r'1ount rkKin1ey
National Park,Alaska ••.•••••••
Seasonal changes in reactions of moose,
observed on the shuttle-tour trips,to the
bus and visitors,t·1ount McKinley National
Park,Alaska ••••••'•••••••••
Behavior before disturbance of moose obs~rved
during shuttle-tour trips,1973~1974,Mount
McKinley Park,Alaska •••••••.•.••
Maximum behavioral reactions of moose observed
during shuttle-tour trips to the bus and
visitors,1973-1974,Mount McKinley National
Park,Alaska •.....••...•.••..
Number of Da1l sheep observed during shuttle-
tour trips,Mount McKinley National Park,
A1as ka . • • • . • . • • • • • • • . • • • •
Sizes of Da1l sheep groups observed during
shuttle-tour trips,Mount McKinley National
Park,Alaska •••••••.••••••••
Average number of Oal1 sheep observed in
study plots,t~ollnt ~1cKinley National Park,
Alaska • • • • • • • • • • • . ••••
PAGE
82
'86
90
93
95
97
106
107
110
xv
Table 28.Reactions of Dall sheep,observed at varying
9istances from the road during shuttle-tour
trips,to the bus and visitors;percent of
animals and groups,1973-1974,Mount McKinley
National Park,Alaska •••••••••••
Table 29.Dall sheep'observed attempting to cross the
road while moving from winter to summer range,
Mount McKinley National Park,Alaska ••••
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Table 30.
Table 31.
Table 32.
Table 33.
rable 34.
Table 35.
Table 36.
Table 37.
Reactions crf Da1l sheep,observed within
200 m of the road during shuttle-tour trips,
to various actions of the bus and visistors;
percent of animals and groups,1973-1974,
Mount rkKinley National Park,Alaska •••••
Behavior before disturbance of Dall sheep
observed during shuttle-tour trips,1973-1974,
Mount McKinley National Park,Alaska •••••
Maximum behavioral reactions of Dall sheep,
observed during shuttle-tour trips,to bus and
visitors,1973-1974,Mount McKinley National
Park,Alaska ............•.•..
Number of brown bears observed during shuttle-
tcrur trips,Mount McKinley National Park,
Alaska .
Distribution of brown bear observations during
shuttle-tour trips by season anrl distance along
road,t10unt rkKinley National Park,Alaska ••.•
Sex and age distributions of brown bears
observed during shuttle-tour trips,Mount
McKinley National Park,Alaska ••.•••
Sizes of brown bear groups observed during
shuttle-tour trips,1973 and 1974,Mount
t~cKinley National Park,Alaska ••••••
Observations of individually recognized brown
bears,Mount McKinley National Park,Alaska
PAGE
112
115
118
124
126
132
133
135
138
140
xvi
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Table 38.Average number of brm'in bears observed in study
plots,Mount McKinley National Park,Alaska 142
Table 39.Reactions of brown bears,observed at varying
distances from the road during shuttle-tour
trips,to the bus and visitors;percent of
animals and groups,1973-1974,~1ount ~kKinley
National Park,Alaska • • • • • • • • • • • •145
Table 40.Reactions of brown bears,ob~erved within 200 m
of the road during shuttle-tour trips,to various
actions of the bus and visitors;percent of
animals and groups,1973-1974,Mount McKinley
National Park,Alaska ••••••••••'..·150
Table 41.Reactions of single brown bears and families,
observed during the shuttle-tour trips,to the
bus and visitors,1973-1974,Mount McKinley
National Park,Alaska • • • • ••• • • • • .155
156
159
172
157
170
168
Behavior before disturbance of brown bears
observed during shuttle-tour trips,1973-1974,
Mount r~cKin1ey National Park,Alaska •••.•
Reactions of brown bears,observed within 200 m
of the road during shuttle-tour trips,to the
bus and visitors as related to behavior before
the disturbance,1973-1974,~1ount r1cKin1ey
National Park,Alaska •.•••••••
Table 42.
Table 43.
Table 44.Maximum behavioral reactions of brown bears,
observed during shuttle-tour trips,to buses
and visitors,1973-1974,~~ount ~1cKinley
National Park,Alaska •••••.•••.••
Table 45.Number of red fox observed during shuttle-tour
.trips,Mount McKinley National Park,Alaska
Table 46.Use of known red fox dens near the r10unt
McKjnley National Park road,1973-1974 ••••
Table 47.Average number of red foxes observed in study
plots,Mount ~·1cKinley National Park,Alaska
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Actions of vehicles and visitors at red fox
dens,1973-1974,Mount ~kKinley National Park,
Al as"ka • • • • • • • • • • • • • • • • • •
Reactions of red foxes at various distances
from the road to vehicles and visitors,1973-
1974,Mount McKinley National Park,Alaska ••
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Table 48 ..
Table 49.
Table 50.Reactions of red foxes,adults and pups at
dens within 200 m of the road,to vehicles
and visitors,1973-1974,Mount McKinley
National Park,Alaska ••••••••••...
PAGE
174
176
178
xviii
Table 51.
Table 52.
Maximum behavioral reaction of red foxes at
dens within 200 m of the road to vehicles and
visitors,as observed during intensive obser-
vations,1973-1974,~4ount McKinley National
Park,Alaska .......•.......•.
Observations of porcupines during shutt1e-
tour trips,1973-1974,Mount McKinley National
Park,Alaska •••••••••••
181
191
Table 53.Numbers of porcupines and snowshoe
observed along the road before and
June 15,1973~1974,Mount McKinley
Park,Alaska •••••••••••
hares
after
National
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Table 54.
Table Al.
Table A2.
Table A3.
Numbers of willow ptarmigan observed during
road census trips in research vehicle,1973-
1974,Mount t~cKinley National Park,Alaska.
Dates of plot observations,~10unt ~kKinley
National Park,Alaska ••••••••••
Actual numbers and minimum densities of
caribou observed during each plot observa-
tion period,first-search,~1ol1nt ~1cKin1ey
National Park,Alaska ••••••••••••••
Actual numbers and minimum densities of
caribou observed during each plot obser-
vation period,total day,~1ount McKinley
National Park,Alaska ••••••••••
199
218
219
220
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Tab le A4.
Tab 1e AS.
Table A6.
Table A7~
Table AB.
Table A9.
Table AlD.
Tab 1e'A11.
Table A12.
Actual numbers and mlnlmum rlensities of moose
observed durin~each plot observation neriod,
first-seal~ch,t10unt t1cl(inley tlational Park,
Alaska .
Actual numbers and minimum densities of
moose observed durinq each plot observa-
tion neriod,total day,t10unt I1cKinlelf
National Park,Alaska •••.••..'••
Actual numbers and minimum densities of
00.11 sheep ohserved durinq each plot obser-
vation period,first-search,1'1ount ~1cKinley
National Park,Alaska .••.•••••••
Actual numbers and minimum densities of
Oall sheep observed durinn each plot obser-
vation period,total day,t10unt '1cKinlev
national Park,Alaska .......••
Actual numbers and minimum densities of
brown bears observed during each plot obser-
vation period,first-search,t~ount t1cKinle.'l
National Park,Alaska •...•••.••••
Actual numbers und minimum densities of
brown bears observed durinq each plot obser-
vation period,total day,i10unt ~1cKinley
National Park,Alaska •..••...•.•.
Actual numbers and minimum densities of
red foxes observec:l during each plot obser-
vation period,first-search,'1ount '1cKinlelf
National Park,Alaska ........•..
.Actual numbers and minimum densities of
red foxes ohserved durinq each plot obser-
vation period,total day,'1ount r~cKinle.v
Natidnal Park,Alaska .......•.•..
Total number of each species of large mammal
observed during each shuttle-tour round-trin
from Riley Carmnround to Eielson Visitor
Center,t10unt '1cKinl~v National Park,Alaska
PAGE
xix
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Table A13.Reactions of caribou,observed at varying
distances from the road during shutt1e~tour
trips,to the bus and visitors;numbers of
animals and groups,1973-1974,Mount McKinley
National Park,Alaska.. • • • • • • • • . •••231
Table A14.Reactions of caribou,observed during shutt1e-
tour trips,to various actions of the bus and
visitors;numbers of animals and groups,1973-
1974,Mount McKinley National Park,Alaska 232
Table A15.Reactions of caribou,observed during shutt1e-
tour trips,to various actions of the ~us and
visitors;percent of animals and groups,1973-
1974,~~ount tkKin1ey National .Park,Alaska 233
Table A16.Reactions of caribou,observed within 200 m
of the road during shutt1e~tour trips,to
various actions of the bus and visitors;
number of animals and groups,1973-1974,
Mount McKinley National Park,Alaska·234
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Table A17.
Table A18.
Table A19.
Table A20.
Reactions of moose,observed at va~ying
distances from the road during shuttle-tour
trips,to the bus and visitors;numbers of
animals and groups,1973-1974,Mount McKinley
National Park,Alaska •.•.•.•••.••
Reactions of moose,observed during shutt1e-
tour trips,to various actions of the bus and _
visitors;number of animals and groups,1973-
1974,Mount McKinley National Park,Alaska
Reactions of moose,observed during shuttle-
tour trips,to various actions of the bus and
visitors;percent of animals and groups,1973-
.1974,t·1ount ~1cKinley National Park,Alaska
Reactions of moose,observed within 200 m
of the road during shuttle-tour trips,to
various actions of the bus and visitors;
number of animals and groups,1973-1974,
Mount McKinley National Park,Alaska
235
236
237
238
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Table A21.
Table A22.
Table A23.
Table A24.
Table A25.
Table A26.
Table A27.
Table A2B.
Reactions of Dall sheep,ohserved at varying
distances from the road during shuttle-tour
trips,to the bus and visitors;number of
animals and groups,1973-1974,Mount NcKinley
National Park,Alaska ••.•••••••
Reactions of Oa11 sheep,observed during
shuttle-tour trips,to various actions of
the bus and visitors;number of animals
and groups,1973-1974,r~ount r·1cKinley
National Park,Alaska • • • • • •
Reactions of Dall sheep,observed during shutt1e-
tour trips,to various actions of the bus and
visitors;percent of animals and groups,1973-
1974,r·1ount rkKinley National Park,Alaska
Reactions of Oall sheep,observed within
200 m of the road during shuttle-tour trips,
to various actions of the bus and visitors;
numbers ·of animals and groups,1973-1974,
Mount rkKinley National Park,Alaska
Reactions of brown bears,ohserved at varying
distances from the road during shuttle-tour
trips,to the bus and visitors;numbers of
animals and groups,1973-1974,Mount McKinley
National Park,Alaska ••••••••••
Reactions of brown bears,observed during
shuttle-tour trips,to various actions of
the bus and visitors;numbers of animals
and groups,1973-1974,r·1ount r1cKinley
National Park,Alaska ••••••••••
Reactions of brown bears,observed during
shuttle-tour trips,to various actions of
the bus and visitors;percent of animals
and groups,1973-1974,Mount McKinley
National Park,Alaska ••••••••••
Reactions of brown bears,observed within
200 m of the road during shuttle-tour trips,
to various actions of the bus and visitors;
numbers of animals and groups,1973-1974,
Mount McKinley National Park,Alaska
PAGE
239
240
241
242
243
244
245
246
xxi
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PAGE
Table A29.Research truck road census trips,dates of
-trips and condition of road access,Mount
McKinley National Park,Alaska • . • . • •247
Table A30.Average daily traffic,traveling east plus west
combined,through Savaqe River Guard Station
1974,~1ount ~1cKinley National Park,Alaska •••248
xxii
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INTRODUCTION
Mount McKinley National Park provides a refuge for a spectacular
array of wildlife native to interior Alaska.The animals are one of
the park's major attractions and most visitors wish to view a variety
of species.The park road traverses much of the prime wildlife habitat
in the park.Park managers hope to optimize visitor use along the road
while minimizing disturbance to the wildlife,thus maintaining the
animals'traditional use of the region and providing visitors the
opportunity to observe wildlife from the road.
Recent dramatic increases in the annual number of visitors to the
park have magnified the potential for disturbance of the wildlife.In
1974,161,000 recreational visitors entered the park (Daniel R.Kuehn,
pers.comm.,1975).Administrators have already taken several regula-
tory measures to minimize human impact on the natural features of the
park.In the early 1950's,the Sable Pass Restricted Wildlife Area was
established.Visitors are required to remain on the road in this area.
In 1972,the National Park Service (NPS)closed th~road to unrestrict-
ed use by private vehicles beyond Savage River (Km 23)and initiated a
free shuttl~bus system.In 1973,the first temporary closed areas
were established aro~nd areas of critical wildlife habitat.No visi-
tors are permitted in these areas.In 1974,a quota system was estab-
lished to limit and distribute backcountry use.
In the past,very little quantitative information has been avail-
able on the effects of various types and levels of human disturbance
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on wildlife.Recently,with the advent of oil development in the far
North,several studies were conducted on the responses of wildlife to
vari ous types of human disturbance,parti cul ar1y responses to ai rcraft
disturbance (Calef and Lortie 1973,Klein 1974,Lenarz 1974,McCourt
and Horstman 1974),noise disturhance (Espmark 1972,McCourt et al.
1974,Memphis State University 1971,P.C.Reynolds 1974),and man-made
structures and seismic line clearinqs (Banfield 1974,Child 1973,Child
and Lent 1973,Klein 1971,McCourt et al.1974,Miller et al.1972).
Only a small body of literature exists on the responses of large mam-
mals to roads and vehicles (Carhaugh et al.1975,Dorrance et al.1975,
Klein 1971,McCullough 1969,Pienaar 1968,Reed et al.1975,Hard et
al.1973),and most available papers are not quantitative.Notes on
the behavioral responses of wild animals to human presence are scat-
tered throughout hundreds of papers on individual species,but there
are only a few comprehensive papers (Altmann 1958a,Bergerud 1974,
Thomson 1972).The effects of human disturbance on the physiology,
energy balance~reproduction,and mortality of mammals are least known.
Geist (19?la,1971b,1975)has reviewed the literature on the effects
of human disturbance on large mammals.He concludes that disturbance
may produce changes in the behavior,distribution,population dynamics,
energy balance,ecology,and physiology of a species.However,he sug-
gests that large mammals can adapt to the presence of man if certain
aspects of their biology are understood and human activities controlled
accordingly to minimize disturbance.
The study reported here was initiated to provide background data
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for use by park managers on human-wildlife interactions along Mount
McKinley National Park road.The study was conducted during the sum-
mers of 1973 and 1974,and focused chiefly on large mammals.The ob-
jectives of the study were five-fold:1)to describe quantitatively
the types and levels of vehicular and visitor activities along the
road,2)to describe quantitatively the behavioral responses of the
large mammals to various human activities,3)to determine the effects
of human activities on the distribution of large mammals near the park
road,4)to provide quantitative baseline data on the present distribu-
tion,relative abundance and population composition of large mammals
along the park road,so that future changes in these parameters and
the effects of future management policies may be assessed,and 5)to
suggest ways to minimize human disturbance to wildlife along the park
road.
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STUDY AREA
Nount r1cKinley National Park covers 7,845 km 2 of interior Alaska,
including parts of the Alaska Range and lands immediately north of the
range (Fig.1).The area is kno\'Jn for its cool,wet summer weather.
Summer precipitation generally comes as frequent,light drizzles.A
general snow cover usually remains into late Nay or early June,and
drifts may last well into the summer.The average annual precipitation
is only about 38 cm (A.Nurie 1944,U.S.Dept.of Commerce 1970).Nay
to September temperature maxima average 10-18°C (U.S.Dept.of Com-
merce 1970).Freezing temperatures may be experienced during any
month.Summer wind speeds vary from 2-9 mlsec (5-20 mph)(U.S.Dept.of
Commerce 1970).From mid-Nay til August there is no complete darkness.
Permafrost occurs through much of the park.
The park road,completed in 1938,extends for 140 km east to west
through the park (Fig.1).It terminates at Kantishna,just outside
the park boundary.All traffic must enter the park,and return,along
the same road.The road 1ies north of,and roughly parall el to,the
Alaska Range.It traverses a low region between the Alaska Range and
the Outer Range.In some areas it reaches high on the south side of
the Outer Range.The elevation along the road varies from 488 m near
Riley Creek Campground at the entrance to the park,to 1,220 m at High-
way Pass (Washburn 1971).The road crosses several major north-flowing
rivers,characterized by broad gravel bars and many changing channels.
Only the first 23 km of the road are paved.It is narrow,with frequent
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MOUNT McKINLEY NATIONAL PARK
Adop/~from:NPMcK-7004,Nov./957,R6V,0I0n./973
--PAVED ROAD --GRAVEL ROADoRANGERSTATIONACAMPGROUND
o 10 20 KILOMETERSe__
Private Vehicle.
Restricted
Beyond This Paint
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Figure 1.Mount McKinley National Park and features of the road,
1973-1974.
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sharp curves,and is designed for slow driving.The road is generally
passable for vehicles at least as far as Eielson Visitor Center (Km
107)from late May to mid-September.It is not kept clear of snow or
open to traffic the rest of the year.In most areas the natural vege-
tation is low and one'can see for long distances.Nost of the work in
the present study was conducted between Riley Creek Campground (Km 0.2)
and Eie1son Visitor Center (Km 107).
The east end of the park,through which the road passes,contains
three main vegetation types (A.Nurie 1944,Viereck and Little 1972)•
Boreal forest occurs to elevations of 762-914 m and somewhat higher
along rivers (A.Murie 1944).Forests are found along the first 33 km
of the road and in several areas where the road descends into major
drainages.White spruce (Picea gLauca)dominates.Aspen (PopuZus
tremuloides)is common in the park headquarters area.The forest
floors are open.Viereck and Little's (1972)"moist tundra"cov~rs
much of the area along the road.In the park,this type is dominated
by low shrubs,including dwarf birch (BetuLa nana),small willows
(SaZix spp.),and blueberries (Vaaainium uliginosum).Taller willows
occur along stream and river beds.Grasses predominate in some areas
and wet regions may contain many sedges.Alpine tundra is found at
higher elevations.Dryas spp.are characteristic of this type.
Heather (Cassiope tetragonal,dwarf willow (Salix reticuZata),and
other low-lying alpine and arctic plants are common.Alpine tundra
occurs along the park road in the Pblychrome Mountain,Highway Pass and
Thorofare Pass areas.Many mountains near the road are topped by talus
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slopes and rocky crags that have very sparse vegetation.
A strip of roadside vegetation,differing from the veyetation
communities through which the road passes,occurs along each side of
the road.The vegetation along the first 2 km of the road includes a
variety of introduced species from a roadside stabilization program
during which a mixture of seeds of exotic plants,including several
grass species,was used.A few of these species have dispersed further
along the road.The vegetation along the rest of the road is generally
composed of native,early successional sp~cies.Willows,grasses,
;
fire\'Jeed (Epi"Lobiwn spp.),horsetail (Equisetwn spp.),and Artemesia
spp.occur frequently along the entire l~ngth of the road.Many other
species occur where conditions are favorable.Many of these plant
species are important food or cover species for various animals.The
strip of roadside vegetation varies in width from a to over 30 m on
both sides of the road;a width of 2-5 m is most conmon.
Reports from the park list 37 species of manmals (A.Murie 1962)and
132 species of birds (A.Murie 1963)occurring in the region.The lar-
ger marrunals include moose (Alces a"Lces),caribou (Rarzgifer tarandus),
Dall sheep (avis dalli),brown bear (Vrsus arctos),wolf (Canis lupus),
red fox (Vu"Lpes vulpes),wolverine (Gulo guZo),and lynx (Lynx cana-
densis)•
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METHODS
I have used several complementary methods to gather data on the
distribution and behavioral responses to human activities of wildlife
along the Nount ~1cKinley National Park road.The study concentrated on
moose,caribou,Da11 sheep,brown bears and red fox.Nany notes were
also taken on wolves,snowshoe hares (Lepus americanus),porcupines
(Erethizon dorsatum),willow ptarmigan (Lagopus Zagopus),long-tailed
jaegers (Stercorarius WYl.gicaudus),and the birds of prey •Because of
the rarity of sightings or limits on time,few data were gathered on
other species.
For one technique,seven study plots,varying in area from 9.8 to
20.7 km2 (Fig.2),were used.Six of the plots were paired (three
pairs),with similar habitat within each pair.One plot of each pair
was placed along the road and the other in the backcountry.The Igloo
Road Plot (15.3 km 2)and Igloo Off-Road Plot (19.2 km 2)both cover
narrow valleys with low but steep,rugged mountains on either side.
Extensive tall willow and some small patches of spruce occur near the
stream beds.At successively higher elevations on the mountains are
found moist (low shrub)tundra,alpine tundra,and talus and rocky
crags.Both plots include large areas of good sheep habitat.The road
passes through the Igloo Road Plot on or near the streambed.The High-
way Road Plot (17.9 km 2)and the Highway Off-Road Plot (12.9 km 2)cover
broader valleys \'li th small amounts of moi st tundra just above spruce
zones and with large areas of alpine tundra.The Stony Road Plot
8
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Figure 2.Location of study plots,1973-1974.
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(12.4 kn;2)and Stony Off-Road Plot (9.8 km 2)include mainly areas of
alpine tundra and wet tundra.Caribou migrate frequently along the
road through the Highway Road Plot and Stony Road Plot.The Sable Road
Plot (20.7 km 2)is in a unique area because of the presence of large
amounts of grasses.
During the summer of 1973,I spent five observation periods in
each plot.In 1974,the Igloo Road Plot and Igloo Off-Road Plot were
visited an additional five times each.The observation periods for
each plot were distributed through the sumn~r,with a similar seasonal
distribution of dates for the members of a pair.Each observation
period began between 0500 and 0800 hours,depending on fog conditions,
and lasted for several hours.The length of observation periods varied
with plot size,topography,vegetation type,weather conditions and
visibility.The dates of each observation period appear in Appendix
Table Al.I walked the same predetermined route during each period
and searched the land within the plot thoroughly with binoculars and
spotting scope,noting the location and behavior of each largemanunal
observed.All human activity in the area was recorded.The observa-
tions were divided into "first search observations",i.e.,all obser-
vations made during the first trip of the day through a plot,and
"daily observations",which include "first search observations"and
observations of new individuals made on the return trip through the
plot.Return trips were less consistent than the first search because
afternoon rains often hampered observations.The numbers of each spe-
cies of large mammal observed per square kilometer in the plots are
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used as minimum densities.
A second technique i nvo 1ved the use of the parlc's shutt1 e bus sys-
tem and the hotel's wildlife tour buses.The wildlife tour buses
differed from the shuttles by having a driver and hostess trained as
tour guides.An observer rode a total of 70 eight-hour round-trips
between Riley Campground (Km 0.2)and Eielson Visitor Center (Km 107)
on these buses;41 trips were taken in 1973 and 29 trips in 19740 Bowdy
Train and Harold Downing were the chief observers on the buses,although
Pat Va1kenburg and I rode as observers on a number of trips.In 1973,
three trips were made each week,one each beginning at 0400 hours,0700
hours,and 1500 hours.In 1974,only two trips were made each week,
one beginning at 0400 hours and one at 1500 hours.The day of the week
for each trip was selected randomly.Twenty-five of the trips were
made on the 0400 hours hotel wildlife tour bus,16 on the 0700 hours
shuttle,and 29 on the 1500 hours shuttle.The times given to identify
the buses are for the beginning of the eight-hour trip.The trips were
taken between May 26 and September 3 both summers (see Appendix Table
A12).Appendix Table A12 shows the type of bus and date of each trip.
During each trip the observer recorded the following information:
1)observer's name,2)date,3)the type and starting time of the bus,
4)the number of people riding the bus,5)the number of buses leaving
at the same time,6)the observer's seat location.For each animal
sighted the following was recorded:1)species,2)individual identi-
fication if known,3)weather and visibility,4)actions of the bus
and people,5}number and .actions of other vehicles an~people in the
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area,6}location,7)time,8)distance of animal from the road,
9)habitat,10)number of animals in a group,11)sex,12)age,
13)color,and 14)behavior.When analyzing the data,each sighting
was considered a "group"even if only one animal was involved.Also,
when the percentage of groups exhibiting some behavior was calculated,
if any animal in a group exhibited the behavior concerned the group
was counted.Some groups were counted more than once if the individ-
uals in the group reacted differently.Thus,percentages for groups
may total more than 100%.
During analysis,each sighting from an entire round-trip of the
bus was considered.Some individual animals could be counted twice on
one trip if they were observed while the bus was traveling out into the
park and resighted during the return.
The reactions of large mammals to the actions of the bus and people
were divided into the following four classes:1)unknown --either trle
reaction was not observed or the behavior of the animal could not unam-
biguously be interpreted as a response to the disturbance,2)no reac-
tion --the animal exhibited no visible reaction to the disturbance,
3)mild reaction --the animal reacted by simply watching the distur-
bance or by moving at a walk less than 10 m,and 4)strong reaction
the animal reacted by moving at a gait faster than a walk or fora
distance greater than 10 m.It must be stressed that this technique
reveals only the instantaneous,visible behavioral reactions shown by
the animals.Nothing is revealed about physiological stress or subtle
behavioral adjustments.Also,nothing is known about the animals that
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react but which are not sighted at all.
A third method involved the use ofa three-quartcr-ton pickup
truck to observe the numbers and distribution of animals near the road.
An "observation trip"consisted of driving the vehicle at approximately
16 km/hr one-way between Riley Campground (Km 0.2)and the Toklat River
(Km 86).The same types of data were recorded as during the shuttle-
tour observation trips.These trips differed from the shuttle-tour
trips in that the speed was slower,and I could stop at any time.Nost
trips began at 0600 hours,and lasted 8-10 hours,depending on the
number and types of sightings.A total of 30 trips was taken in 1973
and 1974 combined.Twelve of the trips were made in late Nay and early
June,just before and after the road was opened to public traffic of
any type.Eighteen of the trips were made in late August and Septem-
ber,just before and after the road was opened to all private vehicles
without limits.These trips provided data on the numbers and distri-
bution of animals along the road during the two periods when visitor
activities were unique by being absent or by having most visitors
driving in private vehicles rather than riding in public buses.The
dates of each trip and the condition of the road access appear in
Appendix Table A29.
Long-term observations of individuals and groups were used as a
fourth technique.Between late July and mid-August 1974,I made in-
tensive observations of caribou between the Toklat River and Eielson
Visitor Center.Several individuals were identified on repeated
sightings by l!sing antler form,body color,size,sex,age and other
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characteristics for identification.During five days,the groups
visible from the road were located five times eachd~y,with time,
location,distance from the road,individual identification,human
activities,and caribou behavior being recorded.In addition,indi-
vidual groups of caribou were observed continuously for periods up
to 7 hours and for a total observation time of 41 hours.During
these periods records were kept on the frequency and types of human
and vehicular activities,the activities and nrovements of each indi-
vidual animal,and the responses of each animal to vehicles and human
activities.
Also in 1974,long-term observations were conducted on denning
red foxes.Four fox families,three inhabiting dens in view of the
road and one family at a den out of view of the road,were observed a
total of 276 hours.Data were recorded on the activities and behavior
of the animals,the frequency and types of human and vehicular activi-
ties,and the reactions of the foxes to these activities.
In addition,many useful observations \'/ere made of wildl ife and
visitor activities during the time spent traveling through and living
in the park in conjunction with this study.
Some data were collected on traffic quantity and quality along the
road.During all work on plots near the road and long-term wildlife
observations,the time,type,and direction of each vehicle was record-
ed.Also,records were kept by NPS of all vehicles passing through the
Savage River Guard Station for the entire s'ummer of 1974.
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Most mileage markers along the road are now missing or incorrect,
and distances cited in visitor manuals vary.During this study a per-
sonal mileage map was developed using the odometer of a three-quarter-
ton pickup truck.
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RESULTS AND DISCUSSION
Visitation and Traffic Levels
Mount McKinley National Park recreational visitation levels have
increased dramatically in recent years (Fig.3).Sharp visitation in-
creases followed developments improving access.In late 1958,the
completion of the Denali Highway connected the park road with the road
system of the state.Before this time access was chiefly via railroad.
In 1972,the new Anchorage-Fairbanks highway (Alaska Route No.3)
opened;by 1973,most of the paving and construction was completed,
making the park an easy day·s drive from either Fairbanks or Anchorage.
Recreational visits increased from 18,000 in 1964 to 161,000 in 1974.
This visitation is concentrated into the summer months,when the park
road is open for vehicle travel.In 1973,98%of all visits occurred
during May through September,with 6%in May,13%in June,39%in July,
27%in August,and 13%in September (U.S.Dept.of Interior 1974).
Winter visitors to the park must travel by skis,snowshoes,or dogsled
if they wish to go further into the park than park headquarters (Km 5).
No snowmobiles are allowed in the park.
During most of the period of this study,1973-1974,private vehi-
cles were not allowed on the park road beyond the Savage River Guard
Station (Km 23)unless the drivers possessed a permit to drive to a
reserved campsite west of the guard station.Only a limited number of
auto-campsites exists.A few special permits were issued to profes-
sional photographers and researchers.There was some through-traffic
16
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197519651955
Year
1945
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1935
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Figure 3.Mount McKinley National Park annual visitation levels,
based on U.S.Department of Interior (1974)report.
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to the Kantishna area west of the park.Most visitors using the road
west of the guard station rode on public buses,either on the free
shuttle bus system or on the hotel's wildlife tour buses.NPS main-
tenance and ranger vehicles also travel the road.
The data collected on traffic moving through the Savage River
Guard Station in 1974 are summarized in Figure 4 (see also Appendix
Table A30).The daily vehicle flow through this station,computed on
a weekly average,increased until the end of June and then remained
between 156 and 176 vehicles until mid-August.This leveling-off of
the traffic flow results from the fact that only a limited number of
private vehicles are allowed,determined by the number of auto-camp-
sites,and that a fairly constant number of public buses travel the
road each day,even though the number of people they carry may vary.
A sharp increase in the number of private vehicles,and thus in the
total number of vehicles,occurred in early September,because the
restriction on private vehicles was removed and a permit was no longer
necessary.This increase occurred despite the fact that the main
tourist season is over by September.The shuttle buses stopped running
in early September.
Figure 5 depicts the hourly"traffic pattern during July 1974 at
the Igloo Canyon-Sable Pass (Km 55 -Km 69)section of the park road.
The graph results from an average of three to seven separate traffic
counts for each hour of the day.Traffic levels ranged from 5 to 12
vehicles/hr from 0700 to 1900 hours,while fewer than 1 vehicle/hr
passed from 2300 to 0600 hours.
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N PS Vehicles
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June July August
For Week Beginning (Month,Day)
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Figure 4.Average daily number of vehicles passinr east plus west
through the Savage River Guard Station in 1974,Mount
McKinley Naticnal Park,Alaska.
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20
Public Buses
2000 2400
"7.5
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0000 0500 1000 1500
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Figure 5.Hourly traffic pattern during July 1974 along the Igloo
Canyon -Sable Pass section of the road,Mount McKinley
National Park,Alaska.
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21
The percentage of various types of vehicles in the traffic flow
varies with the section of road.East of the Savage River Guard Sta-
tion any private vehicle is permitted and,although precise counts were
not conducted,this type of vehicle predominates.Traffic passing
through the guard station in July 1974 consisted of 10%NPS vehicles,
38%public buses,and 52%private vehicles.At the Igloo Canyon-Sable
Pass section of the road during the same period,the composition was
10%NPS vehicles,56%public buses,and 34%private vehicles.The re-
duction in private vehicles reflects the fact that only one campground
exists west of the Igloo-Sable area.
The quantity,quality and diel pattern of the traffic flow along
the park road are all important factors in determining the effects of
traffic and visitors on the wildlife near the road.Relationships be-
tween responses to disturbances and these factors are discussed under
the separate species results which follow.
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Caribou (Rangi[er tarandus)
Population History,Numbers,Composition,Distribution
Very little information is available on the population dynamics,
ecology,or movements of the McKinley caribou herd.Skoog (1968)sum-
marized the existing information,demonstrating a herd history since
1835 of numerous fluctuations in numbers and distribution.Recently,
the herd has decreased dramatically in numbers,with population esti-
mates of 20,000 -30,000 in 1944 (A.Muri.e 1944),12,000 excluding
calves in 1963 (Skoog 1963),3,000 in 1971 (Haber 1972a),and less than
1,000 in 1975 (Willard A.Troyer,pers.conm.1975).The reasons for
the decline are largely unknown,although some evidence exists suggest-
ing emigration (Haber 1972a,Skoog 1968)and possible changes in range
vegetation (Frederick C.Dean,pers.conm.1975).In the last decade,
hunting outside the park and human disturbance inside the park may have
affected the already declining herd,but documentation of these factors
is lacking.The herdls annual movements presently include wintering to
the west and to the north of the park,calving in the foothills of the
Alaska Range in the eastern section of the park,a movement to the
south of the Alaska Range in June,and a migration,especially of cows
and calves,back north and then \'/est,parallel to the park road,in
late June and early July (Hemming 1971).
Caribou were sighted on 90%of the 70 shuttle-tour observation
trips.Table 1 presents the total and average nun~ers of caribou ob-.
served.These.figures exclude large bands with more than 50 animals.
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Table 1.Numbers of caribou observed during shuttle-tour trips,
excluding migrating bands of over 50 anima1s a ,Mount McKinley
National Park,Alaska.
YEAR AVERAGE NUMBER OF AVERAGE NUMBER OF
TOTAL ('IUi'lBER OF CARIBOU OBSERVED "DIFFERENT"CARIBOU
CARIBOU OBSERVED EACH TRIP OBSERVEU EACH TRIP
Animals Groups Animals Groups Animals Groups
1973 808 208 19.7 5.1 17 .5 4.3
1974 371 134 12.8 4.6 10.7 3.9
TOTAL 1179 342 16.8 4.9 14.7 4.1
aReasons for exclusion discussed in text.
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(Note:Bands of over 50 animals occurred along the road for only a
few days during the mass migration each summer.The characteristics
and responses of these large migrating bands were different from,and
are discussed separately from,the small bands that occurred along the
road throughout the summer.)An average of 16.8 caribou in 4.9 groups
was observed each trip.An attempt was made to identify animals that
were sighted twice during one round-trip.If all recognized resight-
ings are excluded,an average of 14.7 "different"caribou in 4.1 groups
was observed each trip.The decrease of the average number of Jldif-
ferent Jl caribou from 17.5 animals per trip in 1973 to 10.7 animals per
trip in 1974 is the result of a peak in caribou numbers during the
first two weeks of June 1973 which did not occur in 1974 (Appendix
Table A12).
The numbers of caribou observed from the various types of buses,
i.e.0400 tour,0700 shuttle,and 1500 shuttle,were compared by pair-
ing trips on the various buses that were separated by no more than two
days.A Wilcoxon Rank Sum test (Wilcoxon and Wilcox 1964)was then
used.For 1973,no significant differences in the numbers of caribou
sighted from the various bus types occurred although the average number
of caribou sighted was higher on the 1500 shuttle than on the 0400
tour.For 1974,the Wilcoxon Rank Sum test showed significantly
(0.005<P<0.Ol)more caribou observed each trip on the 0400 tour than
on the 1500 shuttle.Three factors may be acting to produce these
results:1)~'!eather,2)time of day,and 3)trained guides.~1y rec-
ords of the weather and visibility during the bus trips in 1973 show
24
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that the visibility on the 0400 tour was unlimited or good only 61%of
the time,.\<lhile this value was 91%for the 1500 shuttle.In 1974,
these values were 73%and 76%,respectively.Given similar weather,
more caribou may be observed from the 0400 tour than the 1500 shuttle
because of increased caribou activity during the morning hours.Also,
the tour buses have trained guides who may spot more animals;however,
we found that our trained observers seldom spotted animals not also
observed by someone else on the bus,simply due to.the large number
of people intently watching for wildlife.In some years people on
the 0400 tour may observe fewer animals than people on later buses
because of frequent fog and poor weather in the early mornings.
Large bands of 50-800 animals migrating to the west were observed
in view of the road from July 7 through July 12 in 1973 and from June
29 through July 6 in 1974 (Table 2).Large bands were probably in the
vicinity of the road for a somewhat longer period each year.Small
bands of fewer than 50 caribou occur along the road throughout the
summer.A peak in numbers of caribou in small bands occurred in June
1973,with as many as 146 animals sighted on one shuttle-tour trip.The
number of caribou decreased from mid-August to September to almost zero
both years (Appendix Table A12).
Caribou are distributed in varying frequencies along the length
of the park road because of varying habitat,and,perhaps,because of
traditional use patterns.During the shuttle-tour trips no caribou
were sighted between Km 0 and Savage River (Kin 23).Only 5%of the
caribou occurred between Savage River and Tattler Creek (Km 60),while
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Table 2.Sightings of migrating bands of over 50 carihou from the
road,t~ount tkKinley National Park,Alaska.
DATE OBSERVER NUMBER OF LOCATION
CARIBOU
7/7/73 G.Haber a 800 Polychrome Pass,2000 m S~of road
7/10/73 D.Tracy 320 Highway Pass to Eielson,200-2000 m S.
7/11 /73 of road
7/12/73 B.Train 300 b Eielson,500-900 m S.of road
6/30/74 D.Tracy 251 Km 80.0,near Toklat,300-2000 m S.of
road
6/30/74 D.Tracy 140 Km 78.0,near Toklat,800 m S.of road
7/6/74 c.Downing 132 b Km 79.5,near Toklat,300-400 m S.of
road
7/6/74 D.Tracy 181 Toklat River-Highway Pass,200-500 m
S.of road
apersonal communication.
bObserved during the shuttle-tour trips.
26
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95%occurred between Tattler Creek and Eielson Visitor Center (Km 107).
The same pattern prevailed both summers.Bands tended to move west
during late summer,but caribou were still sighted on Sable Pass (Km
63)in September.
Caribou group sizes,excluding the migrating herds,tended to be
small.Table 3 presents the group size data.Solitary caribou com-
prised 41%of all caribou IIgroupsll,and 12%of all caribou obseryed.
Groups of more than five animals accounted for only 15%of all groups,
but 48%of all animals.The average group size,excluding migrating
herds,was 3.9 animals in 1973.This value declined to 2.8 in 1974.
Group sizes of five or less were observed throughout both summers.In
'1973,most groups larger than five It/ere sighted the first two weeks in
June,but in 1974 the few sightings of groups larger than five were
scattered through the summer.Groups larger than 10 occurred chiefly
west of Polychrome Mountain (Km 74),while smaller groups were sighted
in all areas utilized by caribou along the road •
Table 4 contains the data on the age and sex distributions of the
caribou observed on the shuttle-tour trips.Distance and time available
made it impossible to determine the sex and age of many of the animals.
The relative lack of calves outside the large migrating herds should be
noted.General observations indicate that the migrating herds contain
chiefly cows,calves,yearlings,and,perhaps,young bulls,which are
not easily distinguishable from cows.Two herds that were carefully
counted in 1974 contained a total of 432 animals,including 73 calves,
15 yearlings,26 large bulls,and 318 cows or young bulls.Caribou
~,r-,\-:--1)~'I•.j c-j (t'"J (~r--"'.'----I --I ~(.'1-:-",:.~C'r:-J r-:-J ~i~J r-J\".J \\.L.__,·._,,,J \,J J '.J
Table 3.Sizes of caribou groups observed during shuttle-tour trips,excluding the migrating bands of
over 50 animals a •Mount McKinley National Park.Alaska.
CARIBOU GROUPS ANnlALS
GROUP
SIZE Tota 1 Total
1973 1974 1973-1974 1973 1974 1973~1974
(%of (%of (%of Grand (%of (%of (%of Grand
No.Total)No.Total)No.Total)No.Total)No.Tota 1)No.Total)
1 77 (37)63 (47)140 (41)77 (10)63 (17)140 (12)
2 33 (16 )22 (16 )55 (16 )66 ~8)44 (12)110 (9)
3 30 ~14)18 (13)48 (14 ~90 n~54 (15 )144 (12 )
4 20.10)3 (2)23 ( 7 80 12 (3)92 (8)
5 12 C6)13 (10)25 ~7)60 (7)65 (18 )125 g6~6-10 22 (11 )11 (8)33 10)1li3 (20)75 (20)238
10-50 14 (7)4 (3)18 (5)272 (34)58 (16 )330 (28
TOTAL 208 134 342 808 371 "79
aReasons for exclusion discussed in text.
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Table 4.Sex and age distributions ofcarihou observed during
shuttle-tour trips,~1ount ~1cKin1ey National Park,Alaska.
1973 1974 TOTAL
Number (Percent Number (Percent Number (Percent
Animals Animals)Animals Animals)Animals Animals)
SEX
Male 216 (27)147 (40)363 (31)
Female 154 (l9 )113 (30)267 (23)
Unknown 438 .(54)111 (30)549 (46)
Total 808 371 1179
AGE
Calf 15 (2)8 (2)23 (2)
Yearling 9 (1)13 (4)22 (2)
Adult 459 (57)259 (70)718 (61)
Unknown 325 (40)91 (24)416 (35)
Total 808 371 1179
29
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that remain east of Eielson after the main migration west are predomi-
natelybulls,particularly late in the summer.
During the shuttle-tour observation trips,caribou were observed
in a variety of habitats.Of all caribou sighted outside the migrating
herds,38%occurred in dwarf shrub habitat,24%in alpine tundra areas,
21%on gravel bars and moraines,6%in wet meadows,5%in tall willows
and alder,4?;in tussock tundra,2%on talus slopes,and 1%on snow-
beds.These figures do not necessarily represent the distribution of
habitat use by caribou in the park.The figures are given to show
what habitats caribou along the road were in.
.Human Disturbance
General Avoidance
Using paired plots along the road and in the backcountry,I at-
tempted to determine if caribou avoided the vicinity of the road.
Table 5 summarizes these data.The density data show no evidence that
caribou,in large numbers,avoid watersheds through which the road
passes.In addition,a comparison of the research vehicle observation
trips with the shuttle-tour observation trips showed no changes in
numbers of caribou along the road which could be attributed to the
.opening of the road to bus traffic in the spring or to all vehicles
in the fall.At the time that the road was opened in the fall to
unlimited vehicular use few caribou remained near the road,as they
had already moved toward fall and winter ranges.
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Table 5.Average number of caribou observed in study plots.Mount McKinley National Park.Alaska.
PLOT AVERAGE fJAILY AVERAGE DAILY NO.Of
TOTAL OBSERVATIONS FIRST SEARCH OBSERVATIONS DAYS
ANIMALS
Actual fJens1t 2 Actual Densit2 OBSERVED
Number (per km )Number (per km )OUT OF
FIVE
Animals Groups Animals Groups Animals Groups Animals Groups POSSIBLE
Igloo Road 1973 0.40 0.40 0.03 0.03 0.40 0.40 0.03 0.03
Igloo Road 1974 1.40 1.20 0.09 0.08 1.20 1.00 0.08 0.06 5
Igloo Off-Road 1973 0.40 0.20 0.02 0.01 0.40 0.20 0.02 0.01
Igloo Off-Road 1974 0.20 0.20 0.01 0.01 0.20 0.20 .0.01 0.01
Highway Road 1973 2.20 1.40 0.12 0.08 1.60 1.20 0.09 0.07 4
Highway Off-Road 1973 0 a 0 0 0 a a 0 0
Stony Road 1973 2.00 0.80 0.16 0.06 1.80 0.60 0.14 .0.05 2
Stony Off-Road 1973 2.20 1.40 0.22 0.14 1.60 1.20 0.16 0.12
Sable Road 1973 8.40 4.00 0.41 0.19 6.80 3.20 0.33 0.16 5
w.....
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32
The plot data do indicate that,during the summer,caribou may
use areas near the road more intensively than watersheds with similar
habitat north of the road.Pairing all first-search observation peri-
ods for each set of paired plots (Appendix Table A2)and applying a
matched-pair t-test (Sokal and Rohlf 1969)reveals a significantly
larger number of caribou observed per km 2 in the Igloo Road Plot 1974
(t=4.16,O.Ol<P<O.02)and in the Highway Road Plot 1973 (t=2.78,
O.02<P<O.05)than in the corresponding backcountry plots.Other pairs
showed no significant differences.The unpaired Sabl~Road Plot 1973
had densities of caribou at least twice as great as any of the other
plots and appears to be a favored habitat.These data are consistent
with the fact that the road was built in the same east-west fault
valley utilized as the traditional major caribou migration route.This
distribution pattern increases the potential importance of human dis-
turbance along the park road to the caribou population.No plots were
placed in the higher valleys south of the road;this area may also be
heavily used by caribou.
Baskin (1974)felt that reindeer in eastern USSR prefer to main-
tain a distance of at least one kilometer from man.Klein (1971)
reported that a herd of wild reindeer in Norway abandoned a section of
their range cut off.by a highway and a railroad,after several years
of crossing.He emphasized the possible importance of the cumulative
effects of more than one obstruction.In the present study,road
crossings by single and small bands of caribou were frequently ob-
served,but the large migrating bands moved west parallel to and south
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33
of the road without crossing.
Although the study indicates that caribou are not,in general,
avoiding the area of the park road,the nature of the study provides
little insight on what individual differences may occur.Dorrance et
aZ.(1975),during a telemetric study of white-tailed deer (OdocoiZeus
virginianus),found that one doe with a fawn moved her home range over
a mile from a new source of human disturbance,while other deer in the
area did not change their home ranges.During the intensive caribou
observations in late July and August 1974,I learned to identify cer-
tain individual caribou using characteristics of antler form,size,
coat condition and color,scars,etc.I sighted certain individuals
repeatedly in view of the road,including a young bull sighted over a
period of 12 days,an adult bull for a period of 5 days,and four
single cO\'JS for 6 days,6 days,8 days and 10 days.These are minimum
periods,as the intensive observations covered only a short section of
the road over only a three week period,and it required time to learn
to recognize individuals.These observations show that at least some
individual caribou remain in the vicinity or return to the vicinity of
the road over a period of days or weeks and do not simply move quickly
through the area.Bergerud (1974)reported that several individually
recognizable caribou bulls in Newfoundland exhibited restricted home
ranges during the fly season in July and August •
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34
Reaction Distances
The shuttle-tour data relating the disturbance reactions of cari-
bou to their distance from the disturbance appears in Table 6 and Fig-
ure 6.The percentage of caribou showing visible reactions to buses
and-visitors,and the ratio of strong to mild reactions,decrease with
distance from the road.A G-test (Sokal and Rohlf 1969)performed on
the number of caribou in each known reaction class at 100 m intervals
from the road up to 500 m shO\llS that the pattern is significant
(G=197.2,P<0.005),i.e.there is a relationship between d.istance from
road and reaction.The behavior of 9%of the animals \'Ias classified in
the "unknown"class because of difficulties in interpreting the cause
of some behavior patterns.Animals classified with unknown reactions
were excluded when calculating reaction percentages.Sixty-one percent
of the caribou within 100 m of the road exhibited some visible reaction
to bus-related human activities,41%exhibited strong reactions.Be-
yond 400 m from the road,less than 10%of the caribou gave visible
reactions to road-related disturbances.
Mild reactions were underestimated by the shuttle-tour technique.
Table 7 and Figure 7 compare the distance-reaction data generated from
the shuttle-tour trips and from the intensive observations.Calves
were excluded in this comparison.Their reactions differed from older
animals and the percentage of calf observations \lIas greater during the
intensive observations than during the shuttle-tour observations.
Chi-squared tests show significant differences in the distribution of
reaction strengths (none:mild:strong)for the two techniques for the
l I .--;-r-,'<;i1 .~.~r f)~~~r-r:i i~rr-J CJ J rr--~!--,~,---"1 "--,:-l (jL,.I /,.",L)""'<,,,I,.J'II ..L .,j J J )
Table 6.Reactions of caribou,observed at varying distances from the road during shuttle-tour trips,to the bus and
visitors;percent of animals and groups,1973-1974,Mount McKinley National Park,Alaska.
DISTANCE KNOWN REACTIONS OF CARIBOU TO BUS AND VISITORS TOTAL TOTAL OBSERVATIONS
FROM KNOWN KNOWN AND UNKNOWN
ROAD NONE MILD STRONG REACTIONS REACTIONS
(m)
%Grand %Grand
Percent Percent Percent Percent Percent Percent Number Number Number Number Total Total
Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups
0-25 11 16 32 26 57 58 38 19 42 21 4 6
0-100 39 42 19 25 41 34 217 97 243 104 21 30
0 0 0 46 36 54 64 24 11 26 11 2 3
1-25 28 33 7 11 64 56 14 9 16 11 1 3
26-50 32 44 25 24 43 32 56 25 60 26 5 8
51-75 52 39 39 50 10 11 31 18 33 18 3 5
76-100 50 56 4 11 46 33 92 36 108 40 9 12
101-200 67 77 21 13 12 11 226 71 235 78 20 23
201-300 85 80 10 13 5 7
165 45 168 47 14 14
301-400 83 86 11 7 6 7 105 28'137 32 12 9
401-500 93 90 7 10 0 0 121 21 124 23 11 7
501-750 100 100 0 0 0 a 119 23 125 25 11 7
751-1000 97 92 0 0 3 8 31 12 55 17 5 5
>1000 100 100 0 0 0 a 92 16 92 16 8 5
TOTAL 76 72 12 14 12 15 1076 313 1179 342
Note:"Groups"may total over 100%,since some groups were counted under more than one reaction class when various animals in
the group reacted differently.Raw numbers for animals and groups are presented in Appendix Table Al3.
r
36
N=31....
~
751-1000
Reaction
o None
IZJ Mild
•Strong
....N=1I9
N=121
r-
N=I05
r-
N=165
20 ~~;;f
V'··;o I/t
0-100101-200 201-300301-400 401-500 501-750
Distance from Road (m)
100 r-
80 f-
en N=226
0 ,....
E
c:60 f-
<t-0 N=217-c:40 ~r-IQ)
0'-Q)
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[Figure 6.Reactions of caribou,observed at varying distances from
the road during shuttle-tour trips,to the bus and visitors,
1973-1974,Mount McKinley National Park,Alaska.
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Table 7.Reactions of caribou,at varying distances .from the road,to traffic and visitors,as
observed during shuttle-tour trips and during intensive observations,calves excluded,
1973-1974,Mount McKinley National Park,Alaska.
DISTANCE REACTION OF CARIBOU TO TRAFFIC AND VISITORS TOTAL
FROM ROAD NUI'IBER
(m)NONE f>'IILD STRONG ANH'1ALS
Number (Percent Number (Percent Number (Percent
Animal s Animals)Animals Animals)Animals Animals)
SHUTILE-TOUR
OBSERVATIONS
0-100 83 (39)42 (20)89 (42)214
101-200 151 (67)47 (21 )26 (12)224
201-400 222 (86)23 (9)13 (5 )258
>401 348 (97)8 (2)1 (<1)357
TOTAL 804 120 129 1053
INTENSIVE
OBSERVATIONS
0-100 7 (8)30 (35)48 (56)85
101-200 77 (44)85 (48)15 (8)177
201-400 347 (73)115 (24)14 (3)476
>401 216 (95)12 (5)0 (0)228
TOTAL 647 242 77 966
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_100
o
N=228
N=357
Intensive Observat ions
r-Shuttle-Tour Observations
N=258
~
~
N=224....
!-
Reaction
N=214 0 None
I-IZ]Mild;::.,•Strong
~-,
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20
60
40
100
80
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"-cr 80 N=476
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60 N=85
N=177
40
20
O'-Jo..~_...J.....~L-_---iL..l'...BiL_--_--.L-"'-'I'---
0-100 101-200 201 -400 >400
Distance from Road (m)
Reactions of caribou,excluding calves,to traffic and
visitors;comparison of shuttle-tour observations and
intensive observations,1973-1974,Mount ~1cKinley National
Park,Alaska.
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39
distance classes 0-100 m (;=28.00,P<0.005),101-200 m (X 2=32.86,
P<0.005),and 201-400 m (;=26.41,P<0.005).No significant differ-
ence at the P=0.05 level was found beyond 400 m.The difference
results chiefly from a greater percentage of mild reactions observed
during the intensive observations than during shuttle-tour observa-
tions.During intensive observations,I frequently observed caribou
become alert to a vehicle when it first carne into view at a great
distance,then return to their former activities and give no other
visible reaction when the vehicle reached the closest point to the
animals.Such a reaction often would not be seen by an observer on
a bus.During the intensive observations,no caribou were observed
to note a vehicle in the distance and then purposely move out of view
before the vehicle approached close enough for the animal to be ob-
served from it.Strong reactions were probably not greatly underesti-
mated.Differences in the percentages of strong reactions determined
from the two techniques nay relate to the fact that the intensive
observations include reactions to private and NPS vehicles as well as
to the public buses.A discussion of possible reaction differences
to various types of vehicles appears later.
Quantification of caribou disturbance behavior has been attempted
in other studies,chiefly in relation to caribou reactions to aircraft.
Klein (1974)reported percentages of mild and strong reactions shown
by caribou in response to small airplane overflights that are very
similar,at comparable distances,to the intensive observation data
from my study.He defined a strong reaction as a fast'trot,run,or
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panic reaction.Overflights below 200 ft (60 m)caused 60%strong
reactions and 33~~mil dreacti ons.These percentages were 13%and 49%,
respective1y,~for overflights above 500 ft (150 m).Calef and Lortie
(1973)found caribou responding to aircraft by locomotion only for
overflights at 400 ft (122 m)or below and exhibiting panic reactions
only for overflights at 200 ft (60 m)or lower.However,McCourt et
ale (1974)and McCourt and Horstman (1974)recorded some caribou move-
ment responses to small aircraft even for overflights above 1000 ft
(300 m).They reported thresholds for 50%mild responses (significant
interruption of activity or trotting)and 50%strong responses (running
flight)for airplane overflights at 500 ft (150 m)and 200 ft (60 m),
respectively,and for helicopter overflights at 500 ft (150 m)and
300 ft (90 m),respectively.Geist (1975)provides a critique of the
methods and results of these aircraft disturbance studies.
Kelsall (1957)has suggested 100 yd (90 m)as a IItypical ll flushing
distance for caribou in response to the presence of a man,although he,
de Vos (1960),and Lent (1964)all recognize great variability in
flushing distances,citing instances of a few meters to 1 mi (1.5 km).
In a more detailed analysis,Bergerud (1974)calculated distances
during the spring of 107 ±12 m to alert and 81 ±11 m to flush for
cows without calves,and 181 ±54 m to alert and 164 ±11 m to flush
for bands of cows and calves.
Disturbance Freguencies
A high percentage of caribou within 200 m of the road reacted to
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41
road-related human activities (Table 7).This fact combined with the
frequency of traffic (Figs.4~5)indicates a high degree of interaction
of these-caribou with the traffic.During the intensive observations,
caribou within 200 m of the road averaged 4.5 mild reactions,and 2.5
strong reactions per hour from 0800 through 1700 hours,with a decrease
during night hours.(Note:The diel traffic pattern differs somewhat
in the Toklat River to Eielson Visitor Center section of the road,where
the intensive observations were made,from the Igloo-Sable Pass section,
where the-data for Fig.5 were collected.)Thomson (1973)determined
that wild reindeer activity in Norway was interrupted once every four
hours by disturbing stimuli,half of which were human disturbances.
Non-human disturbances occurred once every eight hours and involved
eagles,ravens,gulls and foxes.Wolves and bears were notably absent
from his study area.In the Tanana Hills,Alaska,Curatolo (1975)
noted a June to October average of one disturbance to caribou from
wolves or bears each 20 hours,and a total non-human disturbance rate
of one disturbance per 11 hours from wolves,bears,eagles,and ravens.
Most of his observations were made during the day-time hours.These
studies indicate that the interruptions of caribou activity caused by
traffic on the park road are much more frequent than natural inter-
ruptions,in the absence of human activities.
Relationship of Reaction to Type of Disturbance
During the shuttle-tour observation trips the bus stopped for 80%
of the caribou sighted,people got out of the bus for 16%of the animals,
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42
and visitors walked off the road toward 4%of the caribou.For caribou
within 200 m of the road these percentages were 88%,15%,and 3%,res-
pectively (Table 8).A chi-squared test on the differences in the fre-
quency pattern of passing-stopping-people getting out for the various
types of buses revealed no significant difference between the 0700
shuttle and the 1.500 shuttle,but did show a significant difference
(X 2=45.76,P<0.005)between the shuttles and the 0400 tour.The tour
stopped for 93%of the caribou sighted while the shuttles stopped for
only 75%.There was no difference between shuttles and tours in the
frequency that people got off the bus.
Table 8 and Figure 8 present the shuttle-tour data relating the
reactions of caribou within 200 m of the road to the type of action of
the bus and visitors (see also AppendiX Table A23).Chi-squared analyses
of the reactions of animals within 200 m of the road showed no signifi-
cant differences in the reaction frequency patterns (none:mild:strong)
for buses passing without stopping and buses stopping,without people
getting out.Significantly different reaction patterns (X 2=8.97,
0.01<P<0.C25)did occur between buses stopping,without people getting
out and with people getting out.The latter action caused an increase
from 26%to 41%in animals exhibiting strong reactions.The sample
size of situations when people got out of the bus and moved off the
road toward caribou within 200 m was too small to warrant statistical
analysis,but this action usually precipitated a strong response from
the caribou (Table 8).
There is some evidence that loud noises increased the percentage
r--:r (-,I~r-m "r,::'J c-J r j [[-""J t~~....--.,r--J ;.......,rJ Jl'""~,.,.,..,"
Table 8.Reactions of caribou,observed within 200 m of the road during shuttle-tour trips,to various actions of the bus and
visitors;percent of animals and groups,1973-1974,Mount McKinley National Park.
ACTION OF BUS KNOWN REACTIONS OF CARIBOU TOTAL TOTAL OBSERVATIONS
AND VISITORS KNOWN KNOWN AN D UNKNOWN
NONE MILD STRONG REACTIONS REACTIONS
%Grand %Grand
Percent Percent Percent Percent Percent Percent Number Number Number Number Total Total
Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups
Pass >15 mph 61 71 21 24 18 6 33 17 36 19 8 10
Pass <15 mph 71 50 24 33 6 17 17 6 19 6 4 3
Stop,people 54 59 22 18 24 24 290 111 314 .122 66 67
remain on
bus,quiet
Stop,people 35 36 24 21 41 43 37 14 39 14 8 8
remai n on
bus,noisy
Stop,people 62 57 7 21 31 28 45 14 49 15 10 8
off bus on
road,qui et
Stop,people 83 67 0 0 17 33 6 3 6 3 2
off bus on
road,noisy
Stop,people 0 0 25 50 75 50 12 2 12 2 2
walk off
road,qui et
Stop,people 0 0 0 0 100 100 3 3 <1 <1
wal k off
road,noisy
TOTAL 53 57 20 20 27 25 '443 168 478 182
Note:"Groups"may total over 100%,since some groups were counted under more than one reaction class when various animals in the
group reacted differently.Raw numbers for animals and groups are presented in Appendix Table A16.
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o None
12I Mild
III Strong
N=327
N=37
Visi tors Remain On Bus
(Quiet)(Noisy)
Bus Stops
N=66
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Off Bus
(All Actions)
44
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Figure 8.Reactions of caribou,observed within 200 m of the road
during shuttle-tour trips,to various actions of the bus
and visitors,1973-1974,t10unt t1cKinley National Park,
Alaska.
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45
and strength of reactions (Table 8).A chi-squared analysis of the
reactions of caribou within 200 m of the road exposed to a bus stopping,
with the people remaining in the bus and quiet,compared with those
exposed to a bus stopping,with people remaining in the bus but making
loud noises,showed significant differences (X 2=6.43,0.025<p<0.05),
although the latter action occurred during only 8%of the sightings,
involving 8%of the animals.On 14 occasions the bus stopped an.d loud
noises were generated by whistling,shouting,honking,banging the side
of the bus,or screeching air brakes,while the people remained on the
bus.These actions increased strong reactions from 24%to 43%.Occasion-
ally,loud noises were accidental,but most often they were purposely
generated to attract the animal·s attention for photographic purposes.
During the 41 hours of intensive observations,I observed visitors
getting out of their vehicles and remaining on the road four times from
196 buses,and four times from 117 private vehicles.People moved off
the road toward caribou twice from the buses and twice from the private
vehicles.In all cases when visitors moved off the road they approached
the animals until they reacted strongly.The sample sizes are too small
to be conclusive,but if one considers the numbers of visitors carried
by each private vehicle and by each bus,on a per visitor basis,dis-
turbances are greater from private vehicles.The actions of visitors
riding public buses may be inhibited by the presence of other visitors
and the bus driver.Some drivers actively attempt to control the ac-
tions of their passengers to minimize disturbances to animals.
The response of an animal to various types of disturbance depends
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partly on the evolutionary development of its senses.Smell is the
most discerning sense in caribou (Bergerud 1974,Kelsall 1968),and they
often depend on smell to warn them of danger (0.J.Murie 1935).Moving
objects are often investigated to receive their scent (Bergerud 1974).
This·behavior was frequently observed when 'visitors got out of their
vehicles,especially when they walked off the road.Sight in caribou
has evolved to perceive movement well,but they seem to have poor sensi-
tivity to color,form,and recognition of motionless objects (Bergerud
1974,Harper 1955,Kelsall 1968).The alert response of caribou to
vehicles at a great distance probably results from the perception of
an unidentified movement.After a vehicle was recognized some caribou
no longer responded to it.Hearing is reported to have little effect
as a danger signal,except when combined with visual stimuli (Bergerud
1974,Ericson 1972,O.J.Murie 1935).Caribou in large groups are sur-
rounded by the noises generated by other caribou (Lent 1964,O.J.Murie
1935),and Lent (1964)considered single caribou to be more responsive to
noise than those in large groups.McCourt et ale (1974)reported that
caribou avoid the intense noise of a simulated gas compressor station
by only a distance of 300 m,and show little behavioral response to the
sound beyond this distance.Ericson (1972)found a semi-domestic rein-
deer herd habituated to the common sounds of shouts,motors,and trains,
but the animals exhibited a learned avoidance response to the sound of a
snowmachine used by men capturing calves.On a few occasions during the
intensive observations,I observed caribou that were out of view of the
road noting traffic sounds and moving to a ridge crest to view the road.
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A similar response to noise is reported by de Vos (1960).
R.elationship of Reaction to Group Size
The shuttle-tour data indicate a relationship between responsive-
ness and group size for caribou within 200 m of the road (Table 9).
Chi-squared tests revealed no significant differences in reactions
between groups of one and two,or between groups of two and three to
five.A comparison of animals in groups of one to five with groups
of six to ten reveals a significant ~ifference (X 2=15.80,P<D.005)in
reaction patterns.The frequency of strong reactions was greater in
the larger groups.Groups larger than ten were too few to analyze.
Both Klein (1974)and McCourt and Horstman (1974)found increases in
the frequency and intensity of reactions of caribou to aircraft over-
flights with increased group size.These results may relate to social
facilitation of reactions.The more sensitive individuals in a group
could stimulate responses in the other members of the group.
The relationship seems to reverse in large herds.Banfield (1954),
de Vos (1960),Harper (1955L Jakimchak et aZ.(1974),Kelsall (1957),
Lent (1964),O.J.Murie (1935),Symington (1965)and Thomson (1973)all
reported de~reased responsiveness of large herds to disturbances.Many
disturbing stimuli may not be perceived by the majority of animals in
a large herd,and disturbed individuals may be calmed by the lack of
response by other herd members (Lent 1964).Child (1973)found caribou
crossing success of a sim1ulated pipeline to decrease with increased
group size and felt that animals in small groups were more investigative.
I'"-C~!~...""~r...'J [~rr----,---,C'J r-J r J [""-""]C"--J ~~r---l rJ .:---J .~.:jLIIoJ,'l'i"",.J J J l J
Table 9.Reactions of caribou,observed during shuttle-tour trips within 200 mof the road,in various size groups to the bus and
visitors,1973-1974,Mount McKinley National Park,Alaska.
GROUP REACTION OF CARI~OU TO BUS ANU VISITORS TOTAL
SIZE
NONE MILD STROi~G
Number {Percent Number {Percent Number (Percent Nurrber (Percent Number (Percent Number (Percent Number Number
Animals Animals)Groups Groups)Animals Animals)Groups Groups)Animals Animals)Groups Groups)An i ma ls Groups
45 (56)45 (56)17 (21)17 (21 )19 (23)19 (23)&1 81
2 40 (64)20 (64)6 (10)3 (10)16 (26)l:l (26)62 31
3-5 85 (62)23 (62)26 (19 )10 (27)26 (19)8 (22)137 37
6-10 52 (48)7 (47)12 (11 )2 (13)45 (41)6 (40)109 15
TOTAL 222 95 61 32 106 41 389 164
Note:Some groups are counted under more than one reaction class if various animals in the group exhibited different types of
reactions.Thus,the nllmber of groups entered under the different reaction classes may total more than the actual total
nllmber of groups given and percents for groups may total over 100%.
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Relationship of Reaction to Age and Sex
Both the shuttle-tour and intensive observations show increased
frequencies'and strengths of reactions for calves compared with adults.
All animals older than calves are considered "adults"because of the
inco'nsistent identification of yearlings.Too few calves were observed
during shuttle-tour trips to provide a sta~istically useful sample.
Table 10 contains the relevant data from the intensive observations,
which involved many observations of only a few individual calves.The
reaction pattern (none:mild:strong)for calves is significantly differ-
ent from those for adults in all distance classes with an adequate num-
ber of calf observations,i.e.101-200 m (X 2=12.29,P<0.005)·and 201-400
m (X 2=28.72,P<0.005).The calves observed were already six to twelve
weeks old and often acted independently of the cows.It was often im-
possible to determine which cow was the dam of a calf,a problem also
noted by Pruitt (1960).The calves often watched vehicles or people,
even at relatively great distances (400-600 m),and followed moving
objects with their eyes.On occasion,calves noted vehicles or visitors
and trotted or ran closer to a cow or cows,even when the cows gave no
visible response.This type of behavior in calves has also been noted
by Calef and Lortie (1973),Espmark (1972),and Klein (1974).Lent
(1964)describes cow-calf disturbance behavior in detail.
The data from this study did not show any differences in reaction
patterns related to sex.In almost half the observations sex was not
determined.'Also,Observations of cows with young calves were limited.
Several authors give evidence that females,especially those with young
r '1 c-:r.:-:J r:-l r:-l r-:J c:-J CJ c-J r--J C--:J h r-J ,.---...,l"J .~c-J .:--J c-JJLJ
Table 10.Reactions of adult and calf caribou,observed during intensive observations,
to vehicles and visitors,1974,Mount McKinley National Park,Alaska.
DISTANCE REACTION OF CARIBOU TO VEHICLES AND VISITORS TOTAL
FRor~ROAD NUt'lBER
(m)NONE MILD STRONG ANIMALS
Number (Percent Number (Percent Number (Percent
Animals Animals)Animals Animals)Animals Animals)
ADULT
0-100 7 (8)30 (35)48 (56)85
101-200 77 (44)85 (48)15 (8)177
201-400 347 (73)115 (24)14 (3)476
>400 216 (95)12 (5 )0 (0)228
Total 647 242 77 966
CALF
0-100 0 (-)1 (-)0 (-)1
101-200 0 (0)11 (65)6 (35)17
.201-400 .10 (29)23 (68)1 (3)34
>400 35 (80)9 (20)0 (0)44
Total 45 44 7 96
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calves t are more wary than males (Bergerud 1974 t de Vos 1960,Dixon
1938 t Ericson 1972 t Espmark 1972 t Lent 1964,Thomson 1972).Ericson
(1972),working with semi-domestic reindeer t found that cows with calves
were the first to respond to disturbances.They initiated movement
away from disturbances,traveled furtherest t and stayed away longest.
Child (1973)reports that nursery bands and bands with female leaders
are the most investigative and most successful at crossing a simulated
pipeline t perhaps because of greater directional motivation.
Along the Mount McKinley National Park road t I observed a few
individual caribou that were particularly unresponsive to human acti~
vities.All these animals were large bulls in the latter half of the
summer.They allowed people to approach them to within 15-25 m without
showing a visible response or giving only a mild response.Dixon (1938)
and O.J.Murie (1935)considered old bulls the least wary of caribou.
Thomson (1973)found rutting wild reindeer bulls least easily disturbed.
Relationship of Reaction to Behavior Before Disturbance
Both shuttle-tour and intensive observations show a relationship
between the reactions of caribou within 200 m of the road to vehicles
and visitors and the animals'behavior before the disturbance (Table
11).There are no significant differences between the reactions of
feeding and lying caribou.However t both techniques indicate a signi-
ficant increase in reactiveness in traveling caribou over both feeding
(for shuttle-tour observations t X2=123.83 t P<0.005;for intensive obser-
vations,X2=28.62,P<O.005)and lying caribou (shuttle-tour,X2=5.24,
Table 11.Reactions of caribou,observed within 200 m of the road during shuttle-tour trips and
during intensive observations,as related to behavior before disturbance,1973-1974,
Nount NcKinley National Park,Alaska.
BEHAVIOR REACTION OF CARIBOU TO VEHICLES AND VISITORS TOTAL
BEFORE NU~1BER
DISTURBANCE NONE NILD .STRONG ANIHALS
Number (Percent Number (Percent Number (Percent
Animals Anima 1s)Animals Animals)Animals Animals)
SHUTTLE-TOUR
OBSERVATIONS
Feed a 145 (82)14 (8)18 (10)177
Feed-walk 14 (52)5 (19)8 (30)27
Lie 53 (72)13 (18)8 (11 )74
Stand b 16 (64)2 (8)7 (28)25
Travel 0 (O)36 (84)7 (16 )43
Total 228 70 48 346
INTENSIVE
OBSERVATIONS
Feed a 43 (38~47 ~4l)24 (21~114
Feed-walk 2 (6 26 74)7 (20 35
Lie 34 (43)36 (46)9 .(11)79
Stand b 2 (29)3 (43)2 (29)7
Travel 3 (11 )'J 4 (15 )20 (74)27
Total 84 116 62 262
a Feeding while traveling at a walk.
b Novement at a walk,trot,or run.
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53
O.05<P<O.1;intensive,X2=39.8l,P<O.005).The "behavior before dis-
turbance"was determined during shuttle-tour trips only when the ani-
ma 15 were sighted before reacti ng to the bus,thus thebehavi or before
disturbance was often unknown for the more sensitive animals.This fact
may explain differences benJeen the results of the two techniques.
During the intensive Observations the behavior before disturbance was
usually known.During these observations I felt that caribou moving
and milling around (classified under IItraveling")within 200 m of the
road were often exhibiting this behavior as a prolonged reaction to
recent prior disturbances,which may explain the increased reactiveness
of these animals.More discussion of this factor appears in the follow-
ing section on activity patterns.Many of the animals lying for long
periods within 200 m of the road may have been habituated individuals.
McCourt et aZ.(1974)reported that feeding and lying caribou bands were
most reactive to aircraft,with lying animals showing the strongest re-
actions.Unlike my study,most of the traveling animals encountered
during the study by McCourt et al.(1974)were in large migrating herds
and had not been subjected to frequently repeated disturbances.Klein
(1974)also found the reactions of caribou to aircraft to be dependent
on the behavior before disturbance;lying groups were slightly more re-
active than feeding groups and traveling groups showed fewer total re-
actions but more strong reactions.
~lationship of Reaction to Time and Season
Data from 'this study did not reveal any reaction differences related
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54
to time of day.The literature also gives little insight into this
factor.However,road crossings by caribou may be more frequent during
times of low traffic frequency.During five days of caribou census
trips between the Toklat River and Eielson Visitor Center,repeated
at 0600,1000,1400,1800 and 2100 hours each day,74%(20 animals,
in 4 groups)of all caribou observed crossing the road were sighted
during the 0600 hour trips and 22%(6 animals,in 2 groups)during
the 2100 hour trips.Bergerud (1974)states that caribou usually
cross the trans-Canada highway early in t~e morning when traffic is
light.These observations could be related either to a preference
for crossing when traffic is light or to a greater amount of traveling
during the night or early morning.
My study included only a short part of the yearly cycle,excluding
late fall,the entire winter and early spring.The shuttle-tour data
for caribou within 200 m of the road do show some changes in reactive
patterns through the summer (Table 12).The data from the summer were
divided into four four-week periods from May 26 to September 15.For
the 1973 shuttle-tour data chi-squared tests showed no significant
differences in reaction patterns for the three four-week periods from
June 23 to September 15.There is a significant difference (X 2=15.22,
P<0.005)between the"caribou reaction patterns exhibited from May 26 to
June 22 as compared to June 23 to September 15;a greater percentage of
the caribou reacted to road-related disturbances the first four weeks
of the tourist season.A similar decline in reactiveness occurred in
1974,but too few caribou were ohserved early in the season to provide
~"r17"';'r--r-:'""l rJ rTIl rr::-J r::-:-)'Cl r--l r-:J L":"J L_,2 r-I r-:'r-J r-1 r-l :-Jlt,'.~L I !
Table 12.Seasonal changes in reactions of caribou,observed within 200 m of the road during
shuttle-tour trips,to the bus and visitors,Nount NcKinley National Park,Alaska.
DATE REACTION UF CARIBOU TO TRAFFIC AND VISITURS TOTAL
PERIOD NUt1BEK
NONE rQILO STkOI~G ANII~ALS
Number (Percent Number (Percent Number (Percent
Animals Animals)Animals Animals)Animals Animals)
1973
r·lay 26-June 22 42 (31)45 (33)50 (36)137
June 23-Sept.15 55 (54)25 (25)21 (21)101
Total 97 70 71 238
1974
f1ay 26-June 22 9 (47)2 (10)8 (42)19
June 23-Sept 15 129 (69)17 (9)40 (22)1<>6
Total 138 19 48 205
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a statistically meaningful sample.A decrease in reactiveness through
the tourist season may be caused by increased habituation in the cari-
bou,an increasing percentage of old bulls among the caribou near the
road,and the onset of the fly season.Kelsall (1957),Klein (1974),
and "Lent (1964)reported a reduced sensitivity to disturbance in cari-
bou in mid-summer when fly harassment was great.At this time the
animals are often in poor conditio~(Zhigunov 1961),and disturbances
area further drain on their already stressed condition.Other sea-
sonal changes in responsiveness reported in the literature include
increased responsiveness to disturbance during winter and during
calving (Lent 1964,Thomson 1972).
Relationship of Reaction to Weather and Habitat
Lent (1964)found differences in disturbance behavior related to
weather conditions,vJith caribou showing the most sensitivity in wind
and storms.Freuchen and Salomonsen (1958)observed the same relation-
ship in reindeer.McCullough (1969)reported increased responses of elk
to vehicles in windy weather,but found no relationship to humidity.
Darling (1937),however,attributed changes in irritability of red deer
(Cervus eZaphus)to changes in humidity and barometric pressure,and
suggested that this resulted from the fact that a warm,moist atmosphere
is a better conductor of scent.
Habitat may affect disturbance behavior.Calef and Lortie (1973)
found caribou in timber to be more reactive than those in the openo
Henshaw (1970)states that caribou show alarm,and hesitate when passing
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57
through thick vegetation.De Bock and Surrendi (1974,cited in Geist
1975)found that caribou approach the Dempster Highway in the Yukon
more readily on the open tundra than in the taiga.The increased wari-
ness in closed vegetation may result from the decreased visibility and
resultant reduced possibilities for the perception of predators.
Effects of Disturbance on Activity Pattern
Table 13 presents a summary of the activities of caribou observed
from the road.The shuttle-tour data are complicated by the fact that
the observer could not always determine the behavior before disturbance.
If traveling animals are more sensitive and less often observed before
reacting,as discussed earlier,this fact ~ay reduce the number of ani-
mals recorded as traveling before disturbance within 200 m of the road.
The shuttle-tour data also include observations from throughout the sum-
mer,including frequent sightings of small migrating bands during June
and July.These migrating bands were usually over 200 m from the road
and thus increased the percentage of animals recorded as traveling in
this distance class.The intensive observations were made on temporary
residents and did not include groups migrating at the time observations
were being made.The intensive observations were also continuous rather
than point observations like the shuttle-tour data.The nature of the
intensive observation data make them more suitable for comparing the
activities of animals at varying distances from the road.A chi-squared
test on these data reveal a significant difference (X 2>500.0,P<0.005)
between the activities of caribou within 200 m of the road and those
"I'~:)
Table 13.Ilehavior oT caribou observed during shuttle-tour trips and intensive ouservations.nUl:1uer of animals eXiliuiting given "initial"
behavior when first sighted during shuttle-tour trips and cariuou-nlinutes of given behavior during intensive ouservations.
1973-1974.Hount NcKinley flational Park.Alaska.
DISTAI,CE l;tlIAVlUR OF t.kkWOU TOTAL
FRON RO,,;)
FEE;)-WALKa TRAVEL b(1:1)FEED LIE STAND OTIiER
SHUTTLE-TOUR flur..ber (Percent Nur.lO~r (Percent J.ur.iber (Percent Ilurlber (P~rcent hur.lo~r (Percent ,lu,"uer (Percent liur.IOE;:r
O&SEi{VATIO:lS Animals Anir.141s)Animals Animals)Animals Anir.:als)Animals Anir.lills)Animals Anin141s)Animals Anir..als)Anir.:als
0-200 1&7 (50)39.(10)74 (20)2S (7)4b (13)0 (il)373
>200 271 (45)1&(3)177 (29)20 (3)122 (20)1 «1)WI
Total 458 57 251 4S 170 901
l/,TEtiSIVE Number c (Percent NudJer (Percent Number (Percent Number (Percent Number (Percent flurrber (Percent Nur.iOer
oSSERVATIONS Minutes Minutes)mnutes Minutes)~linutes Minutes)Minutes Minutes)Mi nutes Minutes)Minutes Minutes)Ninutes
0-200 1099 (39)241 (8)761 (27)37 (1)6&6 (24)3 «I)2027
>200 3969 (56)421 (6)1532 (22)496 (7)605 (8)24 «1 )7047
Total 5068 662 2293 533 1291 27 ~b74
a Feeding while traveling at a walk.
b ~bvement at a walk.trot.or run.
c One caribou exhibiting given behavior for one minute.
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59
beyond this distance.Within 200 m of the road,there is an increase
in traveling at the expense of feeding.This increase seemed to be due
to the frequent disturbances from vehicles and visitors.Thomson (1973)
discovered a severe alteration of the activity pattern of wild reindeer
during a period of intensive hunting;lying and grazing decreased and
traveling increased compared to the same season without hunting.
The general activity patterns indicated by this study (Table 13)
are similar to those reported by other workers.Curatolo (1975)deter-
mined the general activity pattern for caribou in interior Alaska during
post-calving as approximately 53%feeding,27%lying,4%standing,and
17%traveling,and during August dispersal,42%,17%,23%,and 19%,
respectively.(Note:These figures were estimated from a graph in
Curatolo 1975).
Behavior Patterns Related to Disturbance
Several distinct behavior patterns are associated with disturbance
reactions in caribou.Detailed descriptions and discussions of some of
these behaviors appear in Lent (1964)and Pruitt (1960).The maximum
behavioral reactions of caribou observed in my study are summarized in
Table 14.The behavior of caribou II wa tching ll vehicles or people varied
from a casual glance through an alert stance to an alarm pose.Caribou
that were lying and chewing cuds were often observed temporarily ceasing
cud chewing while watching traffic and visitors on the road.Caribou
showing any response to a vehicle or person usually oriented their ears
toward the disturbance,even if they did not look in the direction of
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Table 14.Maximum behavioral reactions of caribou.observed during shuttle-tour and intensive observations.to vehicles and
visitors.1973-1974.110unt NcKin1ey National Park.Alaska.
DISTANCE NAXmUN REACTION OF C1\RIBOU TO VEHICLES AND VISITORS TOTAL
FRON ROAD .NU~BER
(m)NONE ~lATCH ~iALK TROT RUN ANIMALS
NumlJer (Percent Number (Percent Numb~r (Percent Number (Percent Number (Percent
Animals Animals)Animals Animals)Aninl:l1s Animals)Animals Animals)Animals Animals)
SHUTTLE-TOUR
OBSERVATIONS
0-200 235 (53)73 (16)29 (7)57 (13)49 (11 )443
>200 581 (92)35 (6)6 (1)1 «1)10 (2)633
Total 816 108 35 58 59 1076
INTEIlSIVE
OBSERVATIONS
0-200 84 (30)120 (43)8 (3)47 (17)21 (8)280
>200 888 (84)147 (14)3 «1)13 (1)11 (1)1062
Total 972 267 11 60 32 1342
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the disturbance.The alert stance is a "posture in which the animal
stares fixedly with its tail raised and head Upll (Lent 1964).The
alarm pose includes the features of the alert stance,plus the extension
of lI one hind leg out to the side in an exaggeration of the urination
pose"(Lent 1964).Urination was at times observed in conjunction with
the alarm pose during my study.During the intensive observations,the
alarm pose preceded 45%of all strong reactions,and occurred during 1%
of the mild reactions.This posture was generally followed by movement
at either a trot or a run.However,trotting or running from a distur-
bance was not always preceded by the alarm pose.Pruitt (1960)considers
the alarm pose to be a ritualized behavior and it apparently serves as a
social releaser,causing other caribou to become alert,but not to flee
without further stimulus (Lent 1964).
The excitation jump is another disturbance-related social releaser.
An animal exhibiting this behavior "ra ises on its hind legs suddenly,
turns while on its hind legs and usually flees,bounding once or twice
before settling into a trot"(Pruitt 1960).Dugmore (1913),Lent
(1964)and Pruitt (1960)all concluded that this action leaves a scent
from the interdigital gland on the ground at the point of occurrence
and proposed that the scent serves as an olfactory danger signal.Lent
(1964)found that caribou encountering such scent on a trail became
more alert or even avoided the location.Pruitt (1960)and O.J.Murie
(1935)felt that adult bulls gave excitation jumps more frequently than
other groups,but any caribou may show the behavior.During the 41 hours
of intensive observations in my study,excitation jumps were observed on
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eight occasions.A total of 14 animals exhibited this behavior,including
two cows and three calves.On five of the occasions,vehicles or people
were Within 50 m of the animals;on all occasions the disturbing object
was within 300 m.The disturbances causing excitation jumps included:
1)vehicles passing without stopping on three occasions,2)vehicles
stopping without people getting out on three occasions,3)people out
on the road on one occasion,and 4)people off the road approaching the
animals on one occasion.Modified,incomplete jumps were exhibited by
an additional 14 individuals.
Locomotion in caribou,including the walk,trot,and run,is dis-
cussed by Skoog (1968).A walk or trot is generally used during migra-
tions,and a fast trot is used for fleeing over long distances.Because
of their heavy pelage,caribou tire and overheat quickly when running.
O.J.Murie (1935)describes an instance in which a cow and a young
bull died of overheating when caused to run in fright during a capture
attempt.Table 14 shows that 11%of the caribou within 200 m of the
road ran in response to disturbances during shuttle-tour trips,while
13%trotted.Such running and trotting,especially if repeated,affect
the energy budgeting of these animals (Geist 1975).Extended running
could affect their immediate health.Most caribou were observed to run
less than 100 m.Extended running was observed on a few occasions when
caribou,surprised on the road,chose to run down the road for up to
1.5 km in front of approaching vehicles.
Caribou exhibited investigative behavior during some disturbances.
The animals became very alert and approached or circled the disturbing
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object t using a loose t high stepping trot.The animal may have been
attempting to receive ascent from an unidentified object that had been
sighted.During the intensive observations this behavior was observed
on two occasions when caribou approached and circled buses at 3 to 50 mt
and on one occasion when two caribou approached to within 40 m of people
who were off the road.
Sometimes caribou reacting to.a disturbance bunch together.This
behavior was observed on six occasions during the intensive observations.
Bunching has been reported for reindeer and caribou in response to dis-
turbances from aircraft t man t snowmobiles t and wolves (Chi1d and Lent
1973,Chrisler 1956 t Curatolo 1975 t de Vos 1960 t Ericson 1972 t Thomson
1973).On two occasions during the intensive observations calves were
observed to run up to cows.
Temporary leadership may occur during disturbing situations.This
was especially noticeable when bunched groups traveled away from a dis-
turbance or when a group crossed the road.During some road crossings
one animal approached and crossed the road first t followed by the other
members of the groupt often in single file.Thomson (1973)noted tem-
porary leadership during 25%of disturbance reactions of wild reindeer.
Displacement activities t resulting from disturbance-related stress,
have seldom been reported for caribou,perhaps because of the difficulty
of recognizing such behavior.Klein (1974)felt that on a few occasions
"agonistic behavior was stimulated by aircraft disturbances."
Caribou appear to cross the road fairly readilYt but show caution.
During 41 hours of intensive observations,road crossings were observed
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on 16 occasions t involving 63 animals.I observed the details of the
crossings for 53 animals.For all but nine of these animals a vehicle,
often mine,was in view at the time of the crossing.The animals seemed
a\'lal~e of the road as a di sconti nui ty in thei r envi ronment.Cauti on in
crossing was observed even when no vehicles were in view.On four
occasions crossings began as a deliberate approach movement 150-250 m
from the road,and,after the crossing,continued as a deliberate move-
ment away until the animals were 50-200 m from the road.The other
crossings involved shorter movements.Most animals moved at least
40 m away from the road after crossing if vehicles were present.Of
the 53 caribou,the fastest gait used while crossing the road was a walk
for 26%of the animals,a trot for 64%,and a run for 10%.At times
the animals turned and moved along the road once they were on it.This
occurred on three of the 16 above occasions.On two other occasions
caribou exhibiting running behavior stimulated by fly harassment turned
when they came on the road and trotted or ran 20-30 m along the road
before leaving on the same side,without crossing.On another three
occasions,caribou t faced with approaching vehicles while they were on
the road,trotted or ran along the road up to 1.5 km,with the vehicle
followin~,then ran off to the side from which they had come.These
instances were di~rupted crossing attempts.Thus t the road at times
diverted the·direction of movement of animals that came upon it,so
that they moved along the road rather than directly across it.Caribou
harassed by flies,or frightened by approaching vehicles t may perceive the
road as a favorable running surface t smooth and unvegetated.However,
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65
extensive use of the road as a travel route by undisturbed caribou was
not observed.Banfield (1974)and ~lcCourt et aZ.(1974)have found
that caribou in winter often divert their direction of travel to follow
seismic line clearings,perhaps because of favorable snow conditions.
De Bock and Surrendi (1974,cited in Geist 1975)found that caribou
avoided crossing raised sections of the Dempster Highway,Yukon,prefer-
ring to cross where they could see across the road.Child (1973.)found
a similar visual factor important in successful caribou crossings of a
simulated pipeline.
Summary and General Discussion
The park's caribou herd has declined markedly in numbers over
the last decade or more.By 1975,only about 1,000 animals remained
(Willard Troyer,pers.comm.1975).However,small bands of 1-10 cari-
bou were frequently sighted from the road during my study.Caribou
were observed during 90%of the shuttle-tour trips.Large migrating
bands of 50-800 animals were observed along the road for only about a
week in late June or early July each summer.Calves were rarely sight-
ed from the road except in the large migrating bands.
The park road follows the same east-west fault valley that has
been traditionally used as a travel route by caribou during the major
post-calving migration.Also,the plot data showed that significantly
more caribou use this valley as a summer range than valleys north of
the road.Thus,the road passes through an area that is of importance
.to the caribou during the summer and for this reason human disturbance
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along the road could adversely affect the herd.
The plot data indicate that caribou,in general,did not avoid the
vicinity of the park road.Certain individuals may avoid the road,al-
though my data provide little evidence for or against this possibility.
Occasionally,caribou responded to repeated human disturbances by
moving out of view of the road.However,at least during August,some
individuals remained near the road,often in view of it,over a period
of at least two weeks.If such temporary restricted home ranges include
the park road and result in repeated contact with traffic and tourists,
the restricted home ranges could result in habituation to human acti-
vities by individual caribou or,alternatively,in a build-up of stress
in these individuals.Individuals with temperaments favoring habituation
would be most likely to remain for extended periods in the vicinity of
the road.These individuals would be more frequently sighted by tourists
than exceedi ngly wary i ndi vi dua 1s.White et aZ.(1975)noted that res i-
dent caribou at Prudhoe Bay,Alaska appeared to habituate to traffic in
the area,while migratory animals did not.In fkKinley Park avoidance
of the road may be most prevalent in the large migrating bands.They
were observed to approach the road within 200 m only at night when
traffic levels were very low and were never observed to cross the road.
The responses of caribou to human disturbances are influenced by
many variables,including distance from the disturbance,group size,
sex and age of the caribou,past experiences,time,season,and weather.
Although these and other variables have been analyzed separately,they
all interact to determine the response of an individual to a given
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situation at a given time.
Both this study and the literature indicate that caribou generally
react to disturbances only at relatively short distances during the
summer.My results show few strong responses beyond 200 m,and few
visible responses at all beyond 400 m (Table 6).Reported IItypica1 11
flushing distances from a man on foot range from 100 yd (90 m)to 160 m
(Bergerud 1974,Kelsall 1957).Several authors (Harper 1955,O.J.
Murie 1935,Tyrrell 1898)have commented on the IItameness li of caribou.
Bergerud (1974)proposed that caribou disturbance behavior evolved under
the major influences of open habitat,gregariousness,and predation by
wolves.He suggested that flight distances are related to the relative
speeds of caribou and wolves,and that caribou can afford to allow \A/o1ves
to approach in the open until predation intention movements are recognized.
A number of authors (Banfield 1954,Jakimchuk et al.1974,A.Murie 1944)
commented on caribou showing little response to wolves moving 100 m away.
Such behavior may result from a balance of the danger and the advantages
of not expending large amounts of energy unnecessarily.Bergerud (1974)
reported that one caribou herd that was reduced by human hunting to less
than 100 animals after wolves were exterminated from its range has since
developed flushing distances much greater than usual,possibly an adapta-
tion to human hunting in the absence of wolves.
In Mount McKinley National Park,where wolves occur and hunting is
not permitted,short flushing distances could benefit both caribou and
visitors.If visitors learn not to exhibit II pre dation intention movements,"
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such as qUick movements or direct approaches,they can observe caribou
at relatively close range without forcing the animals to abandon areas
with easy visitor access,such as along the park road.
The shuttle-tour buses stopped for 88%of the caribou sighted
within 200 m of the road and people got off the bus for 15%of the ani-
mals.Stopping,without people getting out did not significantly in-
crease the frequency or strengths of responses over passing without
stopping.Producing loud noises or getting out of the bus did signifi-
cantly increase the frequency and strength of responses.The increased
response when loud noises were produced by the bus and visitors may
have resulted from the unexpectedness of the noises,rather than to a
particular sensitivity to noises.Such noises occurred during only 8%
of the bus disturbances.Thomson (1972)reported that sudden,strange
sounds were particularly disturbing to wild reindeer.Both Lent (1964)
and de Vos (1960)noted caribou responding to gunshots 0.5 mi (1 km)
away.Geist (197la)pointed out that animals strive to live in a pre-
dictable environment and that unexpected stimuli are disturbing.
The ability of animals to habituate to various stimuli may depend
on the evolutionary significance of those stimuli.Animals may exhibit
a fixed response to stimuli of great importance in their predator-prey
relationships (Bergerud 1974).Thomson (1972)found that the response
of wild reindeer to various disturbances waned with repetition,but
such habituation did not occur in the case of human scent.Strong re-
sponses to the direct approach of humans may persist because of its
similarity to stimuli from attacking wolves.
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Responsiveness of caribou to disturbances increased with increas-
ing group size for the small bands of 1-10 animals most frequ~ntly
sighted along the road.The literature indicates that responsiveness
is decreased in very large herds (de Vos 1960,Kelsall 1957,Lent 1964,
Thomson 1973).
My data revealed that 6-12 week old calves were more sensitive to
human disturbances than adults.Some old bulls were found to be least
wary.Many authors have found cows with calves most sensitive (Bergerud
1974,de Vos 1960,Lent 1964)and old bulls least sensitive (0.J.~1urie
1935,Thomson 1973)to disturbances.Bergerud (1974)suggested that the
evolutionary fitness of the female relates to the safety and'survival of
her calf and thus to increased wariness,while the fitness of the male
relates to his size,condition,and attentiveness to females during rut.
The male especially must conserve and store energy to be successful
during the rut and to survive the following winter.
The responsiveness of the caribou decreased through the summer.
This decrease may have been associated with habituation and/or the on-
set of the fly season.Lent (1964)and Thomson (1972)found increased
responsiveness of caribou and reindeer to disturbances during winter
and during calving.It is to the advantage of the McKinley caribou
herd that no road-related human disturbances occur during these two
annual critical periods.During winter the McKinley Park road is
closed.Most calving occurs in areas away from the road.However,
non-motorized winter recreation is increasing in the park,along with
motorized (snowmobiles,etc.)recreation outside the park.
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Excitation jumpst a response to disturbance which leaves a scent
on the ground that serves as an olfactory danger signal t occurred fre-
quently.This behavior could cause additional stress t through olfactory
stimuli.t in caribou using areas near the road.
The activity pattern of caribou within 200 m of the road was sig-
nificantly altered t as compared with caribou at greater distances from
the road.Human disturbances increased the amount of time spent moving
about and decreased the time spent feeding.The frequency of activity
interruption caused by human disturbances .for caribou within 200 m of
the road t an average of 4.5 mild reactions and 2.5 strong reactions per
hour from 0800 through 1700 hours t is many times greater than the one
interruption every 8-11 hours from non-human disturbances noted by
Curatolo (1975)and Thomson (1973).What physiological stress this
frequency of disturbance may cause in the caribou is not known.The
alteration of the activity pattern t with less feeding t and the frequent
activity interruptions may adversely affect the energy balance of the
ca ri bou .....li th in 200 m of the road.These effects may remove the s tri p
of caribou range near the road from the amount of range available for
effective utilization.The possible importance of these effects are
magnified b~the data of White et al.(1975)which suggest that adult
caribou are able to obtain net energy for growth and fattening during
only a very short period each year.This period occurs during the sum-
mer.
Geist (1971a t 1971b t 1975)reviewed the effects of disturbance on
caribou and reindeer and presented equations for calculating the energy
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demands of various levels of response to disturbance.Most of the
actual data come from experiences with reindeer reported in Zhigunov
(1961).Diiturbances from herding,snowmobiles,and aircraft at times
resulted in loss in body weight,v/eakened animals with increased sus-
ceptibility to disease,emphysema,absorption of embryos,and desertion
or trampling of calves.
71
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Moose (Alces alces)
'p'p~ulation History,Numbers,Composition,Distribution
No intensive studies have been conducted on McKinley Park's moose
population.Most of the available information has been collected in
conjunction with studies on wolves (Haber 1972a,1972b,A.Murie 1944).
Haber (1972a)estimated the 1970 population of moose in the area north
of the Alaska Range and inside the park,between park headquarters and
the Teklanika-Shushana drainages,at 300 animals.He believes that the
numbers have remained relatively stable for the last decade or more.
In an intensive aerial survey covering the entire park in October and
November 1974,Troyer (1974)counted 624 moose.
Moose were sighted on 91%of the shuttle-tour trips.Table 15
presents the total and average numbers sighted each year.For all 70
trips an average of 5.9 moose in 3.8 groups were observed.The maximum
number observed on one trip was.25 animals.There appears to be a dif-
ference in the number of moose sighted depending on the type of bus,
which is probably related to the time of day.Trips on different bus
types were paired with no more than two days between trips of a pair
and matched~pair t-tests were run.The results showed significantly
more moose sighted on the 0400 tour than on the 1500 shuttle in 1974
(t=2.l1,0.01<P<0.OS).The average number of moose seen on each trip
on the 0400 tour in 1974 was 7.9 animals,while on the 1500 shuttle
the average was 3.7 animals.In 1973,too few trips separated by no
more than two days were taken for statistical comparisons.Time of
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Table 15.Number of moose observed during shuttle-tour trips,~1ount
McKinley National Park,Alaska.
YEAR TOTAL NUMBER OF MOOSE AVERAGE NUMBER OF MOOSE AVERAGE NUMBER OF
OBSERVED OBSERVED EACH TRIP 1101 FFERENT"~100SE
OBSERVED EACH TRIP
Animals Groups Animals Groups Animals Groups
1973 238 153 5.8 3.7 5.5 3.5
1974 174 114 6.0 3.9 5.8 3.8
TOTAL 412 267 5.9 3.8 5.6 3.7
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day seems to influence the number of moose sighted,with the early
morning being the best time.This may be related to the fact that
the moose frequent areas with dense vegetation and are most visible
when active.Weather-related visibility seems less important to the
numbers of moose sighted than to the numbers of caribou sighted,
also probably because they inhabit dense vegetation and can only be
sight~d at relatively short distances,regardless of weather condi-
tions.The only seasonal trend in the numhers of moose observed was
the aggregating of moose for the rut in the Savage River area in
September.
The sex and age distributions of the moose observed appear in
Table 16.The sex of 83%of the moose older than calves was determined,
giving a bull:cow ratio of 41d:10a?The low numbers of bulls sighted
could be related to one or more of three factors:1)an actual skew in
the sex ratio,2)differential distribution and hahitat use by the two
sexes,or 3)more wariness in bulls than cows.Durinq his 1974 aerial
moose survey Troyer (1974)divided the park into a number of survey
areas.For the area of the park north of the Alaska Range between the
eastern boundary and Sable Pass he found a bull:cow ratio of 41d:100~
(n=284).This area includes the section of the road where most of the
shuttle-tour moose observations were made.The identical ratios for
the tVJO studies suggest an actual skew in this area.The number of
bulls is extremely low for an unhunted population,although R.L.
Peterson (1955)found that all reports he received from Alaska showed
sex ratios favoring females.Rausch (1959)reported a bull:covi ratio
1973 1974 TOTAL
Number (Percent Number (Percent Number (Percent
Animal s Animals)Animals Animals)Animals Animals)
SEX
Males 40 (17)40 (23)80 (19 )
Females 108 (45)85 (49)193 (47)
Unknown 90 (38)49 (28)139 (34)
Total 238 174 412
AGE
Single Calves 27 (11 )2 (13)49 (12)
Twin Calves 24 (10)8 (5)32 (8)
Total Calves 51 (21)30 (17)81 (20)
Year1 i ngs "(5)12 (7)23 (6)
Two-Year 01ds 3 (1)2 (1)5 (1)
Adults 154 (65)124 (71 )278 (67)
Unknown 19 (8)6 (3)25 (6)
Total 238 174 412
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Table 16.Sex and aqe distributions of moose observed durinq
shuttle-tour trips,Mount t1cKin1eyNationa1 Park;Alaska.
75
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of830:10o~in a lightly hunted Alaskan population~and ratios of
21-380:10~where more intensive hunting occurred.Reports from Katmai
National Monument~Alaska~for 1969-1972 give ratios of 79-940:100~
(NcKnight 1975).The skewed sex ratio in the eastern part of Hount
McKinley National Park may have been caused partly by the severe win-
ter of 1970-1971.Haber (1972a)felt that bull mortality was particu-
larly high during that winter because of the poor post-rut condition
in which bulls begin the winter.His counts showed bull:cow ratios in
the eastern part of the park of 62o:100~in October 1970 and 36o:100~
in October 1971.It is also possible that the moose in the eastern
section of the park are subjected to some hunting.Winter ranges
extend outside the park (Haber 1972b)in areas easily accessible to
hunters.Troyer's (1974)counts in the park west of the East Fork
River-Sable Pass area~where the moose are infrequently exposed to
hunters~revealed a sex ratio of 1240:100~(n=161).Bulls and cows
seem to be unevenly distributed through the park~at least in late
October and November It/hen Troyer made his surveys.LeResche (1966)
found segregation of the sexes by habitat in a~loose population near
Palmer~Alaska;adult males occupied the hillsides more and valley
floors less than the cows.The.park road is generally low where it
passes through moose habitat~and this factor could affect the sex
ratio of the animals observed from the road.
An age class was determined for 95%of the moose sighted~giving
21%calves,6%yearlings,and 73%two-year olds and older (Table 16).
These figures are probably not suitable for determining a ca1f:cow
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ratio,because cows were often not observed for a sufficient amount of
time to determine the number of calves present.Calves are born in
late May and early June (A.Nurie 1944).Twinning is relatively fre-
quent in this area (my observations,A.Nurie 1944).Moose are rela-
tively long-lived.Bulls reach their prime from 6-10 years of age
(Peterson 1955).Maximum ages are near 18-22 years (Houston 1968,
R.L.Peterson 1955).
Table 17 shows the group sizes for moose observed during the
shuttle-tour trips.Group sizes ranged from 1-8 individuals;77%of
the animals were solitary or in pairs.Except for cow-calf and rutting
associations,moose are a relatively solitary ungulate.Even animals
that appear to be in "groups"may be acting independently,being in
close association only because of the presence of a favored resource
(Denniston 1956,de Vos 1958,Geist 1963,Houston 1974,R.L.Peterson
1955).
Of the moose observed,94%were sighted along the first 60 km of
the road,between Riley Campground and Tattler Creek.Moose occurred
all along this section of road.Further along the road,as far as
Eielson Visitor Center,there are few patches of suitable habitat.Of
the moose sighted,48%were in open spruce-dwarf shrub habitat,16%in
spruce or mixed spruce and Populus woods,18%in dwarf shrub habitat,
14%in tall willow stands,2%on gravel bars,and 1%out on alpine
tundra.
I learned to individually identify only three mOose,one bull with
a deformed antler and one cow with a calf.These animals were observed
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Table 17.Sizes of moose groups observed during shuttle-tour trips,Mount McKinl~y
National Park,Alaska.
MOOSE GROUPS ANIMALS
GROUP
SIZE Total Total
1973 1974 1973-1974 1973 1974 1973-1974
(%of (%of (%of (%of (%of (%of
No.Total)No.Total)No.Total)No.Total)No.Tota 1)No.Total)
1 90 (59)70 (61)160 (60)90 (38)70 (40)160 (39)
2 43 (28)34 (30)77 (29)86 (36)68 (39)154 (38)
3 18 (12 )8 (7)26 (10)54 (23)24 (14 )78 (19 )
4 2 (l)1 (1)3 (l)8 (3)4 (2)12 (3)
8 a (0)1 (1)1 (<1)0 (0)8 (4)8 (2)
TOTAL 153 114 267 238 174 412
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throughout the summer of 1973,all remaining in restricted areas of a
few kilometers along the road.A.t:lurie (1944)noted a cow \'/ith two
calves in the park that remained one entire summer along one 10 km sec-
tion of the road.A.Murie (1961)reported on three bulls that re-
mained together through the summer in an area 3 km across.LeResche
(1974)reviewed the literature on movements and home range size for
North American moose,including Alaska,and concluded that seasonal
home ranges are seldom larger than 5-10 km 2 ,that cows with cal~es
have the smallest seasonal home ranges,and that yearlings and rutting
bulls wander most widely.Individuals often return to the same sea-
sonal ranges in successive years (Houston 1974,LeResche 1974).
Human Disturbance
General Avoidance
Table 18 summarizes the data on moose observations in the plots.
Moose occurred occasionally in all plots,except the Stony Road Plot.
Moose were observed frequently only in the Igloo Road Plot.Individ-
uals appeared to be residing in this plot through the entire sunmer
each year;a cow and calf were sighted several times in 1973,and a
cow and calf,a yearling cow,and an adult bull were frequently sighted
in 1974.Moose apparently were not summer residents in the Igloo Off-
Road Plot.They first appeared in this plot in late August and Septem-
ber.A quantity of shed antlers indicated that wintering moose use
this area.The data do not indicate any large-scale avoidance by moose
of watersheds containing the road.In the Igloo Road Plot,moose were
\\,[,jr1CT':")r:"'""i}r:-n n '\;i /
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Table 18.Average number of moose observed in study plots,Mount McKinley National Park,Alaska.
PLOT AVERAGE DAILY AVERAGE DAILY NO.OF
TOTAL OBSERVATIONS FIRST SEARCH OBSERVATIONS DAYS
ANI1-1ALS
Actual Density Actual Densit~OBSERVED
Number (per km 2)Number (per km )OUT OF
FIVE
Animals Groups Animals Groups Animals Groups Animals Groups POSSIBLE
Igloo Road 1973 1.80 1.20 0.12 0.08 1.40 1.00 0.09 0.06 5
Igloo Road 1974 2.80 2.20 0.18 0.14 2.80 2.20 0.18 0.14 5
Igloo Off-Road 1973 0.60 0.60 0.03 0.03 0.60'0.60 0.03 0.03 2
Igloo Off-Road 1974 0.20 0.20 0.01 0.01 0.20 0.20 0.01 0.01
Hi gh~lay Road 1973 0.20 0.20 0.01 0.01 0.20 0.20 0.01 0.01
Highway Off-Road 1973 0.40 0.40 0.03 0.03 0.40 0.40 0.03 0.03 2
Stony Road 1973 0 0 0 0 0 0 0 0 0
Stony Off-Road 1973 0.20 0.20 0.02 0.02 0.20 0.20 0.02 0.02
Sable Road 1973 0.20 0.20 0.01 0.01 0.20 0.20 0.01 0.01
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often observed noticing passing traffic,but usually they did not move
away long distances.Visitors seldom noticed the moose,v/hich usually
were in dense vegetation.The truck census data showed no reductions
in moose numbers near the road associated with the opening of the road
to public buses in the spring or to all vehicles in the fall.
Reaction Uistances
The distances from the road of moose observed during the shuttle-
tour trips appear in Table 19.Because moose usually occur in areas of
dense vegetation,the moose sighted were distributed closer to the road
than other species,such as caribou,which occur frequently in open
country.Eighty pel"cent of the moose observed "Jere wi tlli n 200 m of the
road.Numbers obsel"ved decreased steadily with distance from the road.
The I"eaction of 15%of the moose was unknown or unintel"preted as
to cause.Of all kno"m reactions,57%of the animals sho\i.Jed no visible
response to the presence of the bus and visitors,18%exhibited a mild
reaction,and 24%exhibited a strong reaction.For animals within
200 m of the road these figures were 50%,20%,and 29%,respectively.
Nild responses may have been underestimated because of the habit of
some moose of slowly drifting away from a source of mild disturbance,
or slowly entering dense cover in response to disturbances.This type
of response may be undetected by an observer on a bus restricted to a
short observation period.This factor is discussed in more detail
later.
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Table 19 and Figure 9 show a steady decline in the percentage of
.moose reacting to vehicles and visitors,and in the ratio of strong to
mild reactions,with increased distance between the animal and the hu-
man activities.A G-test reveals that the relationship between dis-
tance and response is significant (G=57.8,P<0.005).For the statis-
tical test,100 m intervals from 0-500 m were used.Only one group
beyond 400 m from the road was observed to react to human activities
on the road.
McCourt et aZ.(1974)studied the response of a few moose to over-
flights by a Cessna 185.For overflights within 600 ft (180 m)they
recorded 55%nil reactions,38%mild reactions ("significant"interrup-
tion of activity or trotting),and 8%strong reactions (running fl ight)
(n=40).For six moose over 600 ft (180 m)from the aircraft,they
recorded only nil reactions.
The flight distance of moose in response to a man on foot is vari-
able and depends on a variety of factors.However,several "averagell
estimates have been reported by investigators who have become familiar
with moose.For naive moose,reported flight distances include 90-140
yd (80-130 m)(Denniston 1956),100 m (LeResche 1966),150 yd (140 m)
(NcNillan 1954),and 100 m (Stringham 1974).Peterson (1955)found
that some moose fled even at 0.25 mi (400 m)if downwind of a man.
Moose appear to be rather easily habituated to the presence of man if
they are not threatened.Flight distances reported for habituated
moose include 20-50 yd (20-45 m)(Denniston 1956),100 ft (30 m)(de
Vos 1958),25-50 yd (25-45 m)(McMillan 1954),and 30-50 m (Stringham
r
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N=IO
100
>500
.-,.....
N=26
,.....
N=16
c-,.....
N=23
N=59 ,.....
Reaction
c-D None
[2]Mild
N=216 &I Strong,.....
...
~
f-17:1/!;r
1/1/l7~1/V 1/1/~~I Vr;I/D ~Vi"1/,-o
0-100 101-200 201-300 301-400 401-500
Distance from Road (m)
80
20
40
60
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[Figure 9.Reactions of moose,observed at varying distances from the
road d!lrinq shuttle-tour trips,to the bus and visitors,
1973-1974,;'1ount McKinley National Park,Alaska.
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1974).Altmann's (1958a)work illustrates some of the factors affect-
ing flight distances.After several years of work with moose,she
reported the following f1 ight distances:30-70 ft (10-20 m)for CO\'/S
with a newborn calf,150-200 ft (45-60 m)for cows with a heeling calf,
100-120 ft (30-35 m)for bulls in velvet,60-90 ft (20-30 m)for pre-
rut bulls,10-30 ft (5-10 m)for rutting adults,20-60 ft (5-20 m)for
groups in winter,200-300 ft (60-90 m)for animals during a hunting
season,and 15-35 ft (5-10 m)for animals exposed to the familiar ac-
tions of tourists,caretakers,or fishermen.Apparently many of the
mooseA1 tmann observed were habituated to man to some degree.
Relationship of Reaction to Type of Disturbance
Table 20 presents information on the actions of the bus and visi-
tors when moose were sighted.The bus passed,without stopping,14%of
the animals sighted.The bus stopped,\'/ithout people getting out,for
75%of the animals,while the bus stopped,with people getting out,for
11%of the moose sighted.For just the animals within 200 m of the
road,these figures are 12%,75%,and 13%,respectively.The shuttles
tended to pass more animals (22%)than the tours (3%),and people got
off shuttles for more of the animals (14%)than off tours (8%).The
differences in the pattern of activities for tours and shuttles are
significant (X 2=28.45,P<0.005).During the shuttle-tour trips people
were not observed walking off the road,toward moose,but this action
was observed at other times.
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Table 20.Reactions of moose,observed within 200 m of the road during shuttle-tour trips,to various actions of the bus and
visitors;percent of animals and \jroups,1973-1974,"Jount /1cKinley National Park,Alaska.
ACTION OF BUS KNOWN REACTIONS OF MOOSE TOTAL TOTAL OBSERVATIONS
A.'iD VISITORS KrlOWN KNOWN AND UNKrWWN
NONE MILD STRONG REACTIONS REACTIONS
%Grand %Grand
Percent Percent Percent Percent Percent Percent Number Number Number Number Total Total
Animals Groups Animals f1roups Animals Groups"Animals f1roups Animals Groups Animals Groups
Pass >15 r.;ph 70 "71 9 13 22 19 23 21 30 26 9 12
Pass -:15 mph 56 50 11 12 33 38 9 8 10 9 3 4
Stop,people 56 56 21 20 23 26 186 121 221 140 68 69
rer,a in on
bus.quiet
StoP.ceople 5 7 25 33 70 60 20 15 22 16 " 7 7
remain on
GUS.::cisy
Stop,peoole 31 35 23 20 46 45 35 20 41 22 12 1n
off bus on
road.quiet
StoP.people"50 50 50 50 0 0 2 2 3 2
off bus en
road.noisy
StoP.people 0 0 0 0 0 0 n 0 0 0 0 0
walk off
road.quiet
StoP.people 0 0 0 0 0 a n a 0 0 0 0
via 1k off
road."noisy
TOTAL 50 51 20 20 29 30 275 187 327 215
Note:"Groups"may total over 100%,since some qroups were counted under more than one reaction class when various animals in the
group reacted differently.Raw numbers for animals and qroups are presented in Appendix Table .0.20.
87
served moose intensively in national parks,especially Yellowstone and
Grand Teton.Both Altmann and Denniston comment that moose are less
ably relates to a sneak approach,under cover,versus a slow,quiet
approach,but in full view,and with the moose aware of the presence of
~
Altmann (1958a),Denniston (1956),and McMillan (1954)all ob-
disturbed by the noisy,obvious activities of tourists than by the
silent,stealthy approach of an investigator under cover.McMillan,
however,stresses that a slow,quiet approach is less disturbing than
a fast,noisy approach.The difference between the observations prob-
70%.
The reactions of moose are related to the type of disturbance in
Table 20 and Figure 10.Chi-squared tests were used to compare the re-
action patterns exhibited by moose within 200 m of the road in response
to various actions of the bus and visitors.There are no significant
differences in reactions to the bus passing,without stopping,compared
to reactions to the bus stopping,with the people remaining quietly in
the bus.There is a significant difference (X 2=9.08,.P<0.025)in the
reaction patterns of moose exposed to a bus stopping,with the people
quietly remaining on the bus,compared with the bus stopping,with the
people quietly getting off the bus.The latter action increased the
strong reactions from 23%to 46%.There is also a significant difference
(~=23.67,P<O.005)between the reactions of moose exposed to the bus
stopping,with the people remaining quietly on the bus,and those ex-
posed to the bus stopping,with the people remaining on the bus but
making loud noises.Loud noises increased strong reactions from 23%to
f-=
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N=37
N=20
N=186
Reaction
o None
IZJ Mild
•Strong
N=32-
Bus Passes
V0 V7V~V
/V~V
~V V1 V/-v'·v·Vo'--..L....W~:Jl'...;.....L....IV'-lliiiil·'~'L-.&....I'.rl/.....:·iiL·.........II:v~"'---_
Visitors Remain On Bus Off Bus
(Quiet)(Noisy)(All Actions)
Bus Stops
20-
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Figure 10.Reactions of moose,observed within 200 m of the road
dur"ng;LlIttle-t'iur t:"ips,to variolls .:}ctiO~lS of ';;he
bus and visitors,1973-1974,Mount McKinley National
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89
the observer.Moose seem to learn to recognize fami1 iar,harmless human.
activities and often ignore them,while the unfamiliar,threatening,
sneak approach remains disturbing.Moose also seem to recognize indi-
vidual people they see frequently and exhibit fewer responses to them
than to strange people (Altmann 1958a,fvlcNi11an 1954).As with many
mammals,people on foot are more disturbing than vehicles,once a moose
is habituated (Dixon 1938).
The sensitive hearing of moose is important in warning them of
danger (A.Murie 1934,R.L.Peterson 1955,Van Wormer 1972).Moose
are sensitive to the sounds of snapping twigs and rustling brush,often
their first clue of danger in areas of dense cover (McMillan 1954,Van
Wormer 1972).Van Wormer (1972)also reported that moose are sensitive
to high-pitched metallic sounds.The sense of smell is reportedly sen-
sitive in moose (A.Murie 1934,R.L.Peterson 1955).The sense of
sight is poor with respect to the recognition of forms,but sensitive
to movement (Altmann 1958a,A.Murie 1961,Van Wormer 1972).Sight is
apparently least important in detecting danger.
Relationship of Reaction to Sex and Age ------The relationship between sex and age and the reaction to human
activities is given in Table 21.Statistical analyses for animals
within 200 m of the road showed no significant differences between:
1)the reaction patterns of calves versus cows followed by a calf or
2)the reaction patterns of cows without calves versus bulls.However,
there is a significant difference between the combined'pattern of
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Tal>le 21.React ions of va rious sex and a~1e c 1clsses of moose,observed during
shuttle-tour trifJs,to buses and visitors,1973-1974,r·loun~I-icl~inley
Ncltional Park,Alaska.
DISTANCE REACTIOII OF HUOSE TO llUS AIW VISITURS TOTAL
FRON -'WAD lWl1l.iER
(m)NOllE NILD STfWHG A,mlALS
Humber (Percent !lumber (Percent Humber (Percent
Animals Animclls)J\nimals Animals)Anililclls Animals)
CALF
0-100 14 (30)13 (29)19 (41)46
101-200 2 (29)5 (71 )0 (0)7
201-400 I"(71 )1 (14 ) 1 (14 )7:I
>400 4 (67)
2 (33)0 (0)6
Total 25 21 20 66
COH
~IITII CALF
0-100 12 (34)8 (23)1b (43)35
101-200 2 (40)3 (60)0 (0)5
201-400 4 (67)1 (17)1 (17 )6
>400 3 (75)1 (25)0 (0)4
Total 21 13 16 50
CO~I
WITHOUT CALF a
0-100 37 (51 )14 (19)21 (29)72
101-200 18 (72)4 (16)3 (12)25
201-400 15 (88)2 (12 )0 (0)17
>400 3 (100)0 (0)0 (0)3
Total 73 20 24 117
BULL
0-100 24 (52)5 (11 )17 (37)46
101-200 9 (64)3 (21)2 (r4)14
201-400 11 (85)0 (0)2 (15 )13
>400 1 (100 )0 (0)0 (0)1
Total 45 8 21 74
-_.--~.'
a Calf not seen,possible that a few of the~e cows actually had calves.
90
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91
ca 1ves and thei r dams versus the combi ned patterns of cows without
calves and bulls (~=14.28,P<O.005).The reactions of calves and
their dams were interdependent.The calves and dams responded n~re
frequently ana strongly at a given distance,and responded at greater
distances,than the cmJS without calves and bulls (Table 21).A number
of authors have noted the increased wariness of cows with calves and
their greater likelihood of fleeing (Altmann 1958a,Jakimchuk et aZ.
1974,Klein 1974,leResche 1966).
The reactions of cows with calves change with the age of the
calf.For the first 3 days of life,a ca1f 1 s locomotor abilities are
poor (Altmann 1958b)and the cmJ often remains in thick cover,flush-
ing only if approached very closely (Altmann 1958a).The 10colilotor
abilities of calves have some importance in determining the effetts of
disturbance.I have observed very young calves stumble and,on one
occasion,roll ~own a bank,when cows with young calves were surprised
on or near the road by vehicles or off the road by hikers.The poten-
tial for occasional injuries to calves exists.Altmann (1958b)also
reported that young calves occasionally follow humans that have ap-
proached closely and that this behavior may result in an attack on the
person by the cow.When the calf is old enough to follow the cow easily,
the flushing distance becomes very long (Altmann 1958a).The moose calf
depends on the cow for protection,and Geist (1963)argued that calves
are usually not alert to danger.However,we occasionally ooserved
calves responding to buses and visitors by moving closer to the cow,
or hiding behind her,while the cow gave no visible reaction to the
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human activities.Yearlings,only recently deprived of maternal pro-
tection,have been reported ns quite unresponsive (Geist 1963,R.L.
Peterson 19,55).
Relationship of Reaction to Season,Time,Heather,Habitat
The reaction patterns of moose appeared to change through the sum-
mer season (Table 22).For moose within 200 m of the road in 1973,
there were significant differences between the reaction patterns from
Hay 26-July 6 as compared to July 7-August 17 (X 2=13.l4,P<0.005),and
from July 7-August 17 as compared to August 17-September 15 (X2=23.4~,
P<0.005).Trends were similar in 1974,but could not be ana.lyzed sta-
tistically because of the small number of moose observed during the
mid-summer period.The percentage of strong reactions showed little
seasonal change,but mild reactions decreased through the summer.In
1973,but not in 1974,mild reactions i'ncreased again near the end of
the summer.The decreases probably resulted from the habituation of
some individuals through the summer.Animals that only reacted mildly
to human activities early in the summer probably habituated sooner than
those that reacted strongly.It is also probable that less habitua.tion
occurred to the types of stimul i-that generated strong reactions.
Nd1illan (1954),vwrking in Yellowstone,reported that moose were most
sensitive early in the tourist season and habituated as the season pro-
gressed.My study did not include the winter season,although the park
is receiving increasing recreational use in the winter.Altmann
(1958a)reports very short flushing distances for moose in the winter
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Taule 22.Seasonal changes in reactions of 1:loose,observed on the shuttle-tour
trips.to the bus and visitors.Hount Hcl~inley National Park.Alaska •
lJATE:REACTIOH OF HOOSE TO BUS ANU VISITORS .TOTAL
IMil)[K
UISTAilCE rWHE mLl)STRONG All lI-lALS
FROI·j ROAO
(m)Humber (Percent HUlllber (Percent Number (Percent
Animals Animals)Animals Animals)Anilllals Animals)
1973
Nay 26-July 6
0-200 40 (42)25 (26)30 (32)95
>200 21 (75)5 (18)2 (7)28
July 7-August 17
0-200 30 (65)1 (2)15 (33)46
>200 11 (100)0 (0)0 (0)11
Augus t 17 -Sept.15
0-200 10 (31)15 (47)7 (22)32
>200 1 (33)0 (0)2 (67)3
1974
Nay 26-JulY 6
0-200 21 (49)12 (29)10 (23)43
>200 5 (62)3 (38)0 (0)8
July 7-August 17
0-200 12 (67)0 (0)6 (33)18
>200 23 (100)0 (0)0 (0)23
August 17-Sept.15
0-200 25 (61)3 (7)13 (32)41
>200 2 (100)0 (0)0 (0)2
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and relates the fact to reduced vigor and the difficulties of traveling
through deep sno\'/.Altmann also conJlIents on the factor of time in rela-
tion to disturbance.She found moose less sensitive at dusk and dawn
than at other times of the day.
Knowlton (1960)and R.L.Peterson (1955)reported that rr~ose
retreat to areas of dense cover and are less active during windy wea-
ther and storms,when possible \'iarnings of danger,through sounds and
smell,are more difficult to detect.However,de Vos (1958)was unable
to find similar relationships in his study area.Altmann (1958a)and
r~cNillan (1954)found moose more wary in open areas than in areas of
good cover.LeResche (1966)felt that moose are more likely.to flee
precipitously if surprised in heavy cover than if approached in the
open.
Beha vi or Before Di sturbance
The behavior before disturbance for the moose observed on the
shuttle-tour trips appears in Table 23.Increased traveling and de-
creased lying and standing,at least in the open where they are visi-
ble,within 200 m of the road is suggested by the data.
No comparisons could be made between various behaviors before
disturbance and subsequent reactions to disturbances because few non-
feeding moose were observed.~lcNillan (1954)reported that,in Yellow-
stone,lying moose were less easily frightened than feeding animals,
especia lly after they had been bedded down for an hour or more.
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Table 23.Behavior before disturbance of moose observed during shuttle-tour trips,1973-1974,Mount McKinley Park,Alaska.
DISTA:;CE
FR0I1 ROAD
(m)FEEO
GEHAVIOR OF NOuSE BEFOR~DISTUkUA:;CE
FEEO-WALKa LIE STAND
TOTAL
hi-HoER
Ail!HrtLS
NUICber (Percent /lumber (Percent /lumber (Percent flun.ber (Percent i~un;ber (Percent Number (Percent
Anir.lals Anic:als)Animals Animals)Anir.lals Animals)Aninoals Animals)Anillials Anill'oals)"nill.als Animals)
0-200 188 (87)1 «1 )1 «1)5 (2)12 (6)10 (5)217
>200 49 (73)6 (9)3 (4)6 (9)3 (4)0 (0)67
TOTAL 237 (83)7 (2)4 (1)11 (4)15 (5)10 (4)264
a Feeding while traveling at a walk.
b Movement at a walk.trot.or run.
e Alert.drink,interactions.rut activities.
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Behavior Patterns Related to Disturbance
The maximum behavioral responses given by the moose observed dur-
ing the shuttle-tour trips appear in Table 24.Fifty percent of the
animals within 200 m of the road were recorded as showing no visible
response to the bus and visitors.However,this figure probably over-
estimates the true percentage of animals that did not respond.Only
during the second year of the study did I begin to realize the ?ubtle-
ness of some of the responses given by moose.I frequently noted moose
standing completely still;and others browsing slowly along until they
disappeared into cover,not to reappear.Although many of these moose
never looked directly at the bus,they were probably responding to the
presence of the vehicle.A delayed response and freezing stance appear
to be characteristic of the reactions of moose to disturbances (Altmann
1958a,de Vos 1958,Geist 1963,Stringham 1974).
Only 11%of the moose within 200 m of the road were observed to
watch the bus and ~isitors wtthout then moving away_Denniston (1956)
and McMillan (1954)describe the alert posture of moose,which includes
raising the head,holding the ears erect and directed toward the source
of disturbance,and staring intently for several seconds or minutes.If
the animal becomes alarmed it may IIfreeze ll (my observations,Altmann
1958a,Stringham 1974),bring its hind legs together and urinate on
them (Geist 1963,McMillan 1954,A.Murie 1934,R.L.Peterson 1955),
and give a short,harsh bark (de Vos 1958,McMillan 1954).Also,in a
detailed behavior study,Stringham (1971)found that his presence near
moose that appeared habituated but were actually still disturbed
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Table 24.Maximum behavioral reactions of moose observed during shuttle-tour trips to the bus and visitors,1973-1974,
~~unt McKinley National Park.Alaska.
UISTANCE NAXHIUN REACTION OF MOOSE TO BUS AND VISITORS TOTAL
FRON ROAD
(m)NONE WATCH WALK <10 m WALK >10 m TROT.UR RUN
Number (Percent Humber (Percent Number (Percent Number (Percent Number (Percent
Animals Animals)Animals Animals)Animals Animals)AninJals Animals)Animals Animals)
0-200 138 (50)3D (11 ~26 (9)26 (9)55 (20)275
>200 63 (84)6 (8 2 (3)0 (0)4 (5)75
TOTAL 201 36 28 26 59 350
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increased comfort movements from 2-5 per hr to 20-50 per hr,increased
eliminations from 1-2 per hr to 4-8 per hr,and increased the rate of
respiration from 1 every 2-6 sec to 2 per sec.He also noted a decrease
in the length of feeding and rumination bouts.
For moose within 200 m of the road,18%moved away from the bus
and visitors at a walk,and 20%moved away at a trot or run.Flight
is usually delayed for several seconds or minutes after a disturbance
has been detected (my observations,Altmann 1958a,de Vos 1958,Geist
1963),and fleeing animals usually stop frequently to look back (my
observations,Geist 1963,LeResche 1966,R.L.Peterson 1955).Flight
is usually towarps cover (my observations,de Vos 1958,McMillan 1954,
R.~.Peterson 1955).Moose observed from the shuttle-tour buses often
moved slowly away or into cover,while continuing to feed.It was
often difficult to determine if the movement was in response to the
bus and visitors or simply normal movement \'1hile feeding.However,
after some experience,the observers learned to recognize purposeful,
directional movements by some moose that continued to feed after the
bus arrived.Stringham (1974)noted that undisturbed moose moved aim-
lessly over an area several body lengths in radius for an hour or more
while feeding,while disturbed moose,although appearing to feed
"aimlessly",drifted away from the source of disturbance.He found
that if the disturbance continued movement increased and feeding de-
creased until finally the animal stopped feeding and walked or trotted
away.Intense displacement feeding has been reported as conmon in
disturbed moose,before,during,and after flight (Geist 1963,LeResche
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1966).A moose that moves slowly into cover n~y begin trotting rapidly
away after it has entered cover (n~observations,de Vos 1958,R.L.
Peterson 1955).Alternatively,a moose that moves either slowly or
quickly into cover,may stop and stand silently,after it believes it
is hidden (my observations,R.L.Peterson 1955).This type of beha-
vior was often observed in response to the shuttle-tour buses and visi-
tors on the road.McMillan (1954)noted that habituated moose in
Vello\'/stone Park usually moved slowly away from a disturbance,while
naive moose usually trotted or ran away.R.L.Peter~on (1955)felt
that moose seldom ran unless suddenly frightened.It was difficult
for us to determine how far retreating moose moved away fro~the road,
since they were usually quickly out of viev/in dense vegetation.Geist
(1963)reported that moose usually do not flee very far.
McMillan (1954)noted that moose usually rested in cover and pre-
ferred resting areas that provided cover between themselves and the
highway.LeResche (1966)found that moose made good use of islands and
peninsulas of cover when moving out into open areas to feed.
Occasionally,nnose,especially cows with calves,have been known
to attack humans (Altmann 1958a,Denniston 1956).Before attacking,
moose move with a stiff warning.gait,raise the hair on their neck and
back,flatten their ears,retract their lips,and often make a chewing
motion with their mouth (Altmann 1958a,Denniston 1956,Geist 1963).
These are warning actions.If the individual disturbing the moose
leaves,attacks seldom occur (Denniston 1956).We observed no attacks
on vehicles or people using the park road,although threat signals were
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observed on rare occasions.
Moose were frequently observed crossing the road.They occasion-
ally ran on the road for up to 1 km in front of approaching vehicles.
One moose was hit and kill ed by a vehic 1e during the period of my study.
Influence of Roadside Vegetation
In many areas along the road willows grow in a strip along the side
of the road where the drainage water runs off the road.On a few occa-
sions moose were observed feeding on these roadside willows.The wil-
lows may occasionally attract or hold moose near the road.However,
willow is abundant in many areas of the park and feeding on the roadside
willows did not seem to occur frequently.
Summary and General Discussion
In 1970,the moose population in the park east of the Teklanika
River,where the road passes through the most moose habitat,was esti-
mated-at 300 animals (Haber 1972a).During my study,moose were sighted
on 91%of the shuttle-tour trips.The sex ratio of the moose observed
that were older than calves was highly skewed toward females with
410:1009,probably because of an actual skewed sex ratio in this area
of the park (Troyer 1974).Of the moose observed during shuttle-tour
trips 21%were calves.Seventy-seven percent of the animals were soli-
tary or in pairs.Hy observations and the literature (Houston 1974,
LeResche 1974,A.Murie 1944,1961)indicate that many individuals
have restricted seasonal home ranges to which they may return in
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isucceSSlve years.They are also relatively long-lived,reaching maxi-
mum ages of 10-22 years (Houston 1968,R.L.Peterson 1950).These
characteristics undoubtedly result in many of the same individual ani-
mals being observed from the road repeatedly throughout a summer and
in successive years.Repeated contacts with human activities would
allow moose to habituate if the activities were of a type that aid not
reinforce the initial alarm response.If habituation did not occur
individuals could be adversely affected by the frequent disturbances.
In the park,moose were observed congregating near Savage River
during the rut.After a literature review,LeResche (1974)concluded
that the moose population from a large area may concentrate $easonally
in small areas of critical habitat and that the misuse by humans of a
small critical area could seriously affect a large number of moose.
The plot data and the research vehicle road census data did not
reveal any evidence that moose were avoiding the general vicinity of
the road.LJenniston {195tij felt that moose avoided a formerly heavily
used area in Wyoming when road construction began there.Responses of
moose to road construction activities are probably quite different than
their responses to an existing road.In Yellowstone National Park,
McMillan (1954)found that naive noose,frightened from a feeding area
by human visitors,did not return that same day.Habituated moose often
did return the same day.
As in other species,a variety of interacting variables affect
the responses of moose to human activities.Flight distances for moose
are relatively short.For moose sighted during shuttle-tour trips
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within 200 m of the road,50%gave no visible response to the bus and
visitors,20%gave mild responses,and 29%gave strong responses.Mild
responses may have been underestimated due to the subtleness of some
responses.The 50%threshold for strong responses was between 25 and
50 m from the road.Reported "average"flight distances from man
range from 80-140 m for naive moose and from 5-50 m for habituated
moose (Altmann 1958a,Denniston 1956,de Vos 195b,LeResche 1966,
Hd1illan 1954,Stringham 1974).The evolutionary factors affecting
flight distance in moose are not completely clear.Like most large
mammals,they learn quickly and habituate to man in areas where they
are not hunted.They may also often depend on cover to remain unde-
tected,rather than on flight to escape.Their major predators are
wolves,man,and perhaps bears.Nech (1970)found that moose often
successfully protected themselves from wolves by standing and fighting
rather than by fleeing.He also found that if a fleeing moose main-
tained a distance of only 100 yd (90 01)between itself and pursuing
wolves the wolves usually gave up after 10-15 sec.Moose seem to de-
pend on their abil ities to stand and fight,or escape easily,more than
on detecti ng \flO 1ves at a distance.Nech reported that duri ng 120 of
131 wolf attacks on moose that he observed,the moose did not detect
the wolves until they had already begun their stalk or charge.
The shuttle-tour buses stopped for 88%of the moose sighted within
200 m of the road,and people got off the bus for 13%of the animals.
Stopping,without people getting out did not significantly change the
frequency or strengths of responses over passing without stopping.The
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frequency and strength of reactions were significantly increased when
loud noises were produced or people got off the bus.The increase in
strong reactions when loud noises occurred was probably related to the
fact that these noises were unfamiliar or unexpected.Loud noises oc-
curred during only 10%of the sightings,involving 12%of the animals.
Moose are particularly sensitive to sound,because in the dense vegeta-
tion'they inhabit sounds are often the first warning of danger (A.
Murie 1934,R.L.Peterson 1955).However,McMillan found that moose
in Yellowstone habituated to car horns,backfires and other highway
sounds.Geist (1963)observed one bull moose feeding calmly within
100 yd (90 m)of a trail crew using a power saw but noted another bull
jump at the sound of an axe 500-600 yd (460-550 m)away.The differ-
ence was probably related to surprise and habituation.
-'~
Cows with calves were found to be significantly more sensitive to
disturbances than cows without calves,or bulls.Calves were common
along the park road and very young calves with poor locomotor abilities
may be injured if forced to flee during a disturbance.
Some habituation apparently occurred each year as the summer pro-
gressed.Although mild reactions decreased through the summer,strong
reactions did not.
Moose disturbance behavior is characterized by standing without
fleeing,short flights,and the use of cover.Many of the key factors
in the disturbance behavior of moose are probably derived from the
moose-wolf relationship as discussed earlier (Mech 1970).Mild re-
sp'onses were probably underestimated during my study because of the
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subtl e di sturbance behavior of moose.Al tmann (1958a)Conl11ents that
"The popular belief that moose do not 'pay much attention'is errone-
ous.Even when eagerly browsing,grazing or standing still,the moose
will closely observe the disturbance and select the most suitable
moment for sl ipping away.II After becoming more fami 1iar with moose,
I began to real i ze the truth of thi s sta tenient.Because many di stur-
bance studies depend on interpreting the instantaneous visible re-
sponses to a disturbance,the level of sensitivity in moose is probably
often underestimated.
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Dall Sheep (avis daZZi)
Population History,Numbers,Composition,Distribution
Several biologists have studied the distribution,population dy-
namics,habitat utilization,and predator relationships of Mount McKin-
ley National Park's Dall sheep population (Haber 1972a,A.Hurie 1944,
Murphy 1973,Sheldon 1930,Whitten 1975).Estimates based on actual
counts have shown the population fluctuating between 500-3,000 animals
since 1945 (Murphy 1973).During 1973,the populatio~'s size was esti-
mated at 1,600 aninmls (Whitten 1975).At this population level,sheep
were sighted during 100%of the 70 shuttle-tour trips.An average of
34.7 animals in 4.6 groups were observed each trip (Table 25).The
maximum number sighted during one trip was 86 sheep.No seasonal
trends in numbers were evident.There were no significant differences
in the numbers si ghted from the various types of buses.
The group sizes of sheep sighted varied from 1-45 animals (Table
26).The larger groups were generally ewe-lamb bands.
Very few of the sheep sighted during the shuttle-tour trips were
sexed or aged,as Qost of the animals were more than 400 m from the
road and the buses did not stop long enough for the observer to gather
this data.In general,ewe-lamb bands were very common,while bands
of old rams were observed only·occasionally.Many rams move up to the
higher valleys of the Alaska Range for the summer.Bands of sheep are
often segregated by sex alld age (Geist 1971,A.Hurie 1944).Using
ground counts,Whitten (1975)determined the population composition in
McKinley Park for the summer of 1973 to be 44%adult ewes,30%adult
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Table 25.Number of Dall sheep observed during shuttle-tour trips,
Mount McKinley National Park,Alaska.
YEAR TOTAL NUMBER AVERAGE NU~1BER AVERAGE NUMBER OF
OF DALL SHEEP OF DALL SHEEP IIDIFFERENT II DALL SHEEP
OBSERVED OBSERVED EACH TRIP OBSERVED EACH TRIP
Animals Groups Animals Groups Animals
1973 1407 201 34.3 5.0 31.0
1974 1020 123 35.2 4.2 31.8
TOTAL 2427 324 34.7 4.6 31.5
106
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Table 26.Sizes of Da11 sheep groups observed during shuttle-tour trips.Mount McKinley
National Park,Alaska.
DALL SHEEP GROUPS ANUlALS
GROUP SIZE
Total Total
1973 1974 1973-1974 1973 1974 1973-1974
(%of (%of (%of (%of (%of (%of
No.Total)No.Total)No.Tota 1)No.Tota 1)No.Total)No.Total)
1 20 (10)11 (9 )31 (10)20 (1)11 (1 )31 (1)
2-5 83 (41)50 (41)133 (41)274 (19)166 (16)440 (18)
1-10 157 (78)94 (76)251 (77)696 (49)430 (42)1126 (46)
11-20 39 (19)17 (14 )56 (17)574 (41)255 (25)829 (34)
21-30 3 (1 )8 (6)11 (4)71 (5)194 (19 )265 (11)
31-40 2 (1)3 (2)5 (2)67 (5)100 (10)167 (7)
41-50 0 (0)1 (1)1 (<1 )0 (0)41 (4)41 (2)
TOTAL 201 123 324 1407 1020 2427
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rams,6%yearlings,and 20%lambs.The life expectancy of sheep varies
with the vigor and gro\'Jth rate of individuals in a population (Geist
1971c),but animals over 10 years of age are generally considered old.
The oldest sheep found by A.Nurie (1944)in ~1cKin1ey Park were 14
years of age.
Most of the sheep range in the park is east of Eielson Visitor
Center (A.Murie 1944,Murphy 1973).The sheep winter in the Outer
Range and,in mild winters,in a few favorable spots in the Alaska
Range.During the summer some of the sheep remain on the winter range,
while others cross the east-west valley through which the road passes
and sunmer in the Alaska Range.Movements between winter and summer
ranges generally occur in June,although there is great variability in
timing (A.Murie 1944).While working with Stone's sheep (avis dalZi
stonei)in Canada,Geist (1971c)concluded that individual sheep show
high fidelity to their seasonal home ranges,that home ranges and mi-
gration routes are transmitted by tradition from old to young animals,
and that the social system minimizes dispersal of young.Geist stressed
the patchiness of suitable sheep habitat,with patches connected by
traditional migration routes.Murphy (1973)pointed out that the range
in McKinley is more continuous than in the area \'/here Geist worked.
However,there is evidence of range fidelity for individual sheep in,
and just outside,the park (my observations,Heimer et aZ.1972,Whit-
ten 1975).
Although sheep range is visible in the distance along most of the
road from Ri1e,y Campground to Eielson Visitor Center,the road passes
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within a few hundred meters of suitable sheep habitat in only a few
places.During the shuttle-tour trips,4%of the sheep sighted were
between Riley Campground (Km 0.2)and Km 51,just before Igloo Canyon,
65%were sighted on Igloo,Cathedral and Sable mountains (Km 51-70),
and 30%were sighted between the East Fork River and Eielson Visitor
Center (Km 70-107).Both A.Murie (1944)and Murphy (1973)found some
of the highest summer densities of sheep in the park in the IgloD-
Cathedral-Sable mountains area.Most of the sheep sighted during the
shuttle-tour trips were high on mountainsides,near escape terrain,
unless they were crossing from one mountain to another.Of the sheep
sighted,47%were on talus slopes,23%were in alpine tundra.20%were
in mixed alpine tundra and talus,5%were in cliffs,10%were in dwarf
shrub-open spruce areas,and 1%or less were in each habitat type of
spruce woods,Populus stands,tall willow,dwarf shrub,and gravel
bars.
The timing of lambing in the park varies yearly,but late ~lay and
early June is the most common time (Dixon 1938,A.Murie 1944).Rut-
ting usually peaks in November (A.Murie 1944,Sheldon 1930).
Human Disturbance
General Avoidance
Sheep were observed in all the study plots except the Stony pair
(Table 27).The Igloo pair of plots contaiQed the highest densities.
The average minimum density of sheep in the Igloo Off-Road Plot was
considerably higher than in the Igloo Road Plot each summer,but a
109
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Table 27.Average number of Dall sheep observed in study plots,Mount McKinley National Park)Alaska.
PLOT AVERAGE nAILY AVERAGE nAILY NO.OF
TOTAL OBSERVATIONS FIRST SEARCH OBSERVATIONS DAYS
ANiMALS
Actual nensit~Actual nensit1 OBSERVED
Number (per km )Number (per km )OUT OF
FIVE
Animals Groups Animals Groups Animals Groups Animals Groups POSSIBLE
Igloo Road 1973 32.80 4.40 2.14 0.29 28.20 3.40 1.84 0.22 5
Igloo Road 1974 17.40 3.20 1.14 0.21 15.69 1.05 1.02 0.21 5
Igloo Off-Road 1973 92.00 6.40 4.79 0.33 56.80 4.80 2.95 0.25 5
Igloo Off-Road 1974 45.00 4.80 2.38 0.25 45.20 4.60 2.35 0.24 5
Highway Road 1973 -4.00 0.80 0.22 0.04 '4.00 0.80 '0.22 0.04 3
Highway Off-Road 1973 3.60 0.60 0.28 0.05 '3.60 0.60 0.28 0.05 2
Stony Road 1973 0 0 0 0 0 0 0 0 0
Stony Off-Road 1973 0 0 0 0 0 0 0 0 0
Sable Road,1973 6.60 1.00 0.32 0.05 6.20 0.80 0.30 0.04 3
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matched-pair t-test on all the first search observation periods (Appen-
dix A6)showed no significant difference at the p=0.05 level.The
Igloo Road Plot receives frequent off-road visitor use.On five occa-
sions during my 10 plot observation periods I observed visitors climb-
ing the mountainside to approach bands of sheep.However,the sheep
appeared quite habituated to this activity,often allowing humans to
approach closer than 100 m,and did not move long distances away unless
a photographer insisted on approaching closer and closer until the
sheep moved a long distance away or moved onto the cliffs.
Most of the road runs below the areas of prime sheep habitat,so
that close contact between sheep and human activities along the road
is infrequent.However,on Polychrome Mountain the road cuts through
the sheep range.Despite the presence of the road,some sheep still
use ~his area,as discussed below.In the past there has been more
use of this area by sheep (A.Murie 1944),but the level of use may be
related to overall population size,density and range factors,as well
as to the presence of the road.
Reaction Distances
Table 28 and Figure 11 show the distances from the road of sheep
sighted during shuttle-tour trips and the reaction patterns of sheep at
varying distances from the road.Only 6%of the animals sighted were
within 200 m of the road.Most of the prime sheep habitat visible from
the road is over 400 m away.Only under two situations were sheep
generally within 200 m of the road:1)when crossing the road during
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N=236N=189-N=15 N=115 r--N=311
r-r-Reaction
-0 None
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II Strong
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0-100 101-200 201-300 301-400 401-500 501-750
Distance from Road (m)
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Figure 11.Reactions of Dall sheep,observed at varying distances
from the road during shuttle-tour trips,to the bus and
visitors,1973-1974,Mount McKinley National Park,Alaska.
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movements from summer range to winter range,and 2)when using the
range on Polychrome t10untain through which the road passes.
Sheep crossing lowland areas between patches of mountainous habi-
tat are very wary,probably because of their increased susceptibility
to predators in the lowland areas (A.Murie 1944).Thus t sheep in this
category are alert and cautious,even before they detect human activi-
ties along the road t and may be expected to be highly sensitive.Eight
crossing attempts \'/ere observed during my study.The dates,times,
locations,and numbers of animals involved appear in Table 29.Most
crossings that occurred when the road is open to traffic take place in
late Nay and June.There were several points along the road where
crossings occurred.Of these eight groups,two groups crossed the road
when no vehicles or people were in view on the road t two groups had
crossed the road and continued across the valley before any vehicles
arrived,one and part of another group crossed cautiously even though
vehicles and people were nearbYt and two and part of another group
turned back and did not cross when vehicles approached during attempted
crossings.The two groups that turned back both ran over 1 km back up
the mountainside from \'/hich they had come before stopping.Thus,the
road does inhibit the movements of some sheep and would affect range
utilization if repeated crossing attempts were thwarted.However,
other individuals exhibited a degree of habituation to crossing in the
presence of vehicles and people.
Both years of n~study,sheep used the Polychrome area,where the
road climbs up into sheep habitat,from ihe time the road was opened
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Table 29.Dall sheep observed attempting to cross the road \'/hile
moving from winter to sununer range,Nount r'1cKinley National
Park,Alaska.
DATE TIME LOCATION NUlvlBER OF CROSSING
ALONG ROAD SHEEP SUCCESSFUL
(km)
1\1ay 24,1973 2220 27.4 7 No
June 3,1973 0800 74.3 4 .Yes
June 16,1973 1245 86.9 2 Yes
June 23,1973 0455 27.4 1 No
June 27,1973 1345 57.1 3 2 Yes,1 No
July 17,1973 1205 14.5 17 Yes
June 20,1974 1000 57.1 2 Yes
July 30,1974 0710 61.2 3 Yes
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until mid-June.In 1974,three full-curl rams remained in the area
into July.It is not known how long in the winter and spring,before
116
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the road opens,this area is used.Both years at least 15 different
sheep were sighted in this area,including rams of all ages,ewes,and
yearl i ngs,but no 1ambs.These sheep frequently res ted and fed wi thi n
50 m of the road,often exhibiting only mild responses to vehicles and
to people taking photographs from 20-50 m away.
The strength of reactions and the percentage of sheep showing
visible responses to buses and ~isitors on the road decrease with
increasing distances between the sheep and the road (Table 28 and Fig.
11).A G-test shows a significant (G=20l.6,P<0.005)relationship be-
tween response and distance for 100 m distance classes from 0-500 m.
Of all the sheep sighted from the buses,only 5%exhibited visible
responses,but for sheep within 200 m of the road,50%of the animals
showed some visible response and 32%gave a strong response.No strong
responses were noted at distance beyond 400 m.
Lenarz (1974)subjected 154 groups of Dall sheep in the Brooks
Range to helicopter flights at diagonal distances of 300-500 ft
(90-l50 m)and reported that 15%of the groups did not move,49%
moved at a walk,and 36%ran away.Dall sheep are apparently capable
of detecting,and may run from,aircraft up to 0.5-2 mi (1-3 km)away
(Andersen 1971,Jakimchuk et al.1974,Linderman 1972).However,
under favorable circumstances,especially after a few days of habitu-
ation,a man on foot in plain view can often approach sheep closely
without causing them to flee.Dixon (1938)and Jones et ale (1963)
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found that they caul d observe some bands of Da 11 sheep from only 150 ft
(50 m).Andersen (1971)reports that he was able to observe Dall rams
from only 50 ft (15 m),and ewes with older lambs from 200 ft (60 m).
Welles and Welles (1961)often watched bighorns (avis cW1adensis)from
100 ft (30 m)and some habituated indivi~uals could be approached to
within 6 ft (2 m)on foot or in a vehicle.Several factors discussed
below affect flight distances for sheep.
Relationship of Reaction to Type of Disturbance
For 25%of the sheep sighted the bus passed without stopping;for
49%of the sheep the bus stopped,with the people remaining on the bus;
and for 21%of the sheep the bus stopped and people got out.For ani-
mals within 200 m of the road,these figures were 14%,72%,and 14%,
respectively.
For sheep within 200 m of the road,the pe~~entage of strong reac-
tions increased between passing and stopping and between stopping and
people getting out (Table 30 and Fig. 12).However,the sample sizes
are insufficient for statistical analysis.There was a significant
difference (X 2=45.59,P<O.OOl)be~Jeen the reactions to the bus stop-
ping,with the people remaining quietly on.the bus,and the bus stop-
ping,with the people remaining on the bus but making loud noises.
All the sheep within 200 m of the road reacted strongly to sudden loud
noises.McCourt et al.(1974)found that Dall sheep avoided the vici-
nity of a noise simulator producing the sounds of a gas compressor
station.However,P.C.Reynolds (1974)observed little response by
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(Quiet)(Noisy)
Bus Stops
Off Bus
(All'Actions)
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Figure 12.Reactions of Da11 sheep,observed within 200 m of the
road during shuttle-tour trips,to various actions of
the bus and visitors,1973-1974,Mount McKinley National
Park,Alaska.
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sheep at a lick to the same noise simulator 0.25 mi (0.40 km)away.
She felt that the sheep were habituated to noises because of a high
level of aircraft activity in the area in the past.
Many researchers working with wild sheep have found that the best
way for a man on foot to approach the animals is to remain in full
view,move slowly,stop often,and not approach directly (Andersen
1971,Couey 1950,Dixon 1938,Geist 1971c,Jones et aLe 1963,Welles
and Welles (950).Sheep flee quickly from a person trying to stalk
close under cover.In their relationships to predators,sheep appar-
ently feel secure if they are near escape terrain and can see the move-
ments of a potential predator.A.Murie (1944)found that wolves were
usually successful in catching sheep in mountainous terrain only if
they could surprise them at.~lose range.He observed that once sheep
sighted a \volf in the distance,they would sometimes move around to
keep the predator in view,or flee if they lost sight of it.He also
observed sheep showing little concern towards wolves resting in full
view only 200 ft (60 01)away.
The direction of approach may also be important.A man approach-
ing from above often causes sheep to flee at a relatively long distance
(Couey 1950,A.Murie (944).The sheep may pass close to a man in
their attempt to flee upward (Smith (954).However,Geist (197lc)pre-
ferred approaching sheep from above so that the wind was favorable.
During my study,sheep were occasionally sighted on slopes below the
road.Often these sheep ran up to the road,across it,and on upwards,
when a vehicl~approached.Some habituated individuals were not
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frightened by vehicles or photographers above them.A.Murie (1944)
noted that one successful hunting technique for wolves is to get above
a band of s'heep and drive them down towards the lowlands.This danger
may explain the wariness of sheep towards humans or other animals above
them.
Sight seems to be the major sense used by sheep to warn them of
danger (Geist 1971c,A.Murie 1961,Welles and Welles 1961).Welles
and Welles (1961)reported that bighorns can spot a moving person up
to 1.5 mi (2.4 km)away.Geist (1971c)found that Stone's sheep could
detect moving objects in obscure terrain much better than he could,
but felt that their power to resolve lines and objects is not good.He
noted sheep staring at animals over 400 yd (370 m)away for 10-20 sec
even if it was only another sheep.The importance of smell to sheep
is unclear.A.Murie (1944)concluded that scent is little used as a
danger signal,perhaps because of the ever changing and complex air
currents in sheep habitat.Geist (1971c)reported that under favorable
conditions sheep can smell a man at 350 yd (321 m).Little is known
about hearing in sheep but their abilHies appear to equal man's
(Geist 1971c,Welles and Welles 1961).
Relationship of Reaction to Sex and Age
Because so few of the sheep in my study were sexed and aged the
effects of these factors cannot be isolated.Other workers have found
that in unhunted or lightly hunted populations old rams are the least
wary and easiest to approach (Andersen 1971,Dixon 1938,Lenarz 1974,
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122
Smith 1954).This characteristic may be reversed in hunted popula-
tions,where rams may be most wary (Jones et al.1963',/YlcCann 1956).
Ewes with 1ambs are particul arly sensitive to disturbances (Jones et
al.1963,A.Murie 1944,Smith 1954).Welles and Welles (1961)found
that a bighorn ewe,that was so habituated she ignored hun~n observers
only 6 ft (2 01)away,fled from the same observers at 200 yd (180 01)
after she gave bi rth to a 1amb.l~hen the 1amb was two and a half
months old,the ewe was still more \'/ary than before the lamb was born.
It should be noted that during my study,lambs \'Jere almost never ob-
served within 200 m of the road.
Relationship of Reaction to Season,Weather,Habitat and Experience
Smith (1954)reported that the bighorns he worked with were more
wary during the summer than during the winter.Jones et al.(1963)
found Dall sheep to be more wary and difficult to approach during pe-
riods of rain,fog,or snow that caused low visiblity.Since sheep
depend on sight to detect and follow the movements of predators,re-
duced visibility nIDy cause increased wariness.
A.~lurie (1944)found sheep to be very wary when crossing valleys
or feeding far from cliffs.Several authors (Andersen 1971,Price
1972,Welles and Welles 1961)reported increased wariness at springs
and mineral licks,\'/hich are often located low on the slopes,away from
escape terrain.As already discussed,sheep observed during n~study
within 200 m of the road were often attempting to cross the valley
through which the road runs and were particularly cautious,alert and
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ready tb flee.uixon (1938),Geist (l971c),Jones et al.(1963),and
A.Nurie (1944)all conillent on the exteme \'iariness of wild sheep
crossing 10.w country,where the risk of predation is greater.They
may spend several hours or days surveyi ng the va 11 ey from an overlook-
ing.ridge before crossing (Geist 1971c,A.~Iurie 1944).During the
actual crossing they often move in compact bands and do much trotting
and running (Dixon 1938,Jones et al.1963,A.Nurie 1944).
~Je11es and Welles (1961)found that the responses to humans of
bighorns in Death Valley depended on the past experiences of individ-
uals and on the leadership of the band.A confident leader appeared
to calm other individuals in a group.One band they observed became
much n~re wary after the loss of its old,habituated leader.
Behavior Before Disturbance
Table 31 contains the data on the behavior before disturbance of
the sheep observed during the shuttle-tour trips.The pattern of
behaviors for sheep within 200 m of the road is significantly different
(X 2=b.b2,P<0.005)from the pattern for sheep more than 200 m from the
road.The major difference is the greater percentage of traveling
animals within 200 m of the roa~,which reflects the animals attempting
to cross the valley.
No analysis could be made on the relationship of behavior before
disturbance to reaction because of the small number of sheep observed
within 200 m of the road.
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Table 31.Behavior before disturbance of Dal1 sheep observed during shuttle-tour trips,
1973-1974,Mount McKinley National Park,Alaska.
DISTANCE BEHAVIOR OF DALL SHEEP BEFORE DISTURBANCE TOTAL
FROM ROAD NUMBER
(m)FEED LIE STAND TRAVEL a AN HlALS
Number (Percent Number (Percent Number (Percent Number (Percent
Animals Animals)Animals Animals)Animals Animals)Animals Animals)
0-200 58 (51)32 (28)0 (0)23 (20)113
>200 1639 (77)299 (14 )76 (4)105 (5 )2119
TOTAL 1697 (76)331 (15)76 (3)128 (6)2232
,a Movement at a walk,trot,run.
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Behavior Patterns Related to Disturbance
Table 32 shows the maximum behavioral reactions exhibited by sheep
toward the buses and visitors.Of the sheep within 200 m of the road,
50%showed no visible reaction,14%watched the bus and visitors,14%
walked away,2%trotted away,and 20%ran away.
Geist (1971c)described the "attention"and "alarm"postures of
wild sheep.These postures may act to draw the attention of others in
a group.Stamping and blowing may accompany the alarm posture (Geist
1971c,Jones et aZ.1963,Welles and Welles 1961).Flight is usually
uphill and often towards escape terrain (my observations,Andersen
1971,Geist 1971c,Jones et aZ.1963,A.Murie 1944,Smith 1954,Welles
and Welles 1961).Such uphill flight occasionally causes sheep in the
park to dash across the road close to approaching vehicles.During
flight,bunching and temporary leadership often occur (my observations,
Dixon 1938,Geist 1971c,A.Murie 1944,Welles and Welles 1961).
Summary and General Discussion
During 1973,the park's Da11 sheep population was estimated at
1,600 aninals.Sheep were sighted during 100%of the shuttle-tour
trips.Groups sizes varied from 1-45 animals.Ewe-lamb bands were
most common.Sixty-five percent of the sheep observed \'/ere sighted
along the 10 km stretch of road near Igloo,Cathedral and Sable
mounta ins.
The road runs below most of the area of prime sheep habitat,so
the sheep are not in close contact with traffic.However,on Igloo
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Table 32.Maximum behaviorai reactions of Call sheep.observed during shuttle-tour trips,to bus and visitors.1973-1974,
Mount '~Kinley National Park.Alaska.
DISTANCE fLl<WIUN REACilCillS OF oru SHEEP TO GUS A;;u VISITORS TuTAL
FRO:·!RO';U ;,iJh.:$tR
(m)NOI.E \lATCH IIALK <10 m wALK >10 m TkOT kUN AJdMALS
Nuriber (Percent !lumber (Percent !lumber (Percent Ilur.llJer (Percent Number (Percent liur.llJer (Percent
AniIMls Animals)Animal s Animals)Animals Anir.:als)Animals Aniloi<11s)Animals Anilii<11s)Anirii<11s Anir.:als)
0-200 65 (50)18 (14)6 (5)11 (9)3 (2)lei (2U)129
>200 2140 (98)35 (2)1 «1)1 «1 )1 «I)9 «1)21 !:a
TOTAL 2205 53 7 12 5 35 £317
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and Cathedral mountains many people get out of vehicles and climb
toward the sheep,which appear quite habituated.The plot data did
not reveal any large-scale range abandonment stimulated by road-related
activities.A few sheep still use the range by the road on Polychrome
Mountain where the road runs up into sheep habitat.Use of this area
by sheep has been much greater in the past (A.~lurie 1944).Geist
(1971)noted short and long-term abandonment of seasonal home ranges
by Stone's sheep caused by hunting.Geist pointed out that hikers in
an area where fear of man is reinforced by hunting may increase the
frequency and seasonal duration of disturbances to sheep and help sti-
mulate range abandonment.Lindern~n (1972)felt that Dall sheep on
Table Hountain in the Brooks Range may have abandoned part of their
summer range because of frequent aircraft disturbances.NcCourt et
aZ.(1974)found that Dall sheep temporarily abandoned the range within
1 mi (1.5 km)of their study area when helicopter activities became
frequent and when a sound simulator,reproducing the sounds of a gas
compressor station,was operating.Welles and Welles (1961)concluded
that bighorns will not continue to usc a desert spring once human
dwellings are established there.
Only 6%of the sheep sighted during shuttle-tour trips were within
200 m of the road.Host of these animals were either attempting to
cross the road while crossing between ranges,or were on Polychrome
Mountain where the road runs through sheep habitat.Qf eight crossing
,--
attempts observed,two groups and part of another turned back and did
not cross because of disturbances from vehicles and people.During
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spring movements from wi nter to SUlliner range many sheep attempt to
cross the road after it is opened to traffic.Disturbances of the
sheep attempting to cross the road may be detrimental.
The buses stopped for 865~of the sheep within 200 m of the road,
and people got out of the buses for 14%of the animals.The frequency
. .
and strengths of reactions increased as the bus stopped and as people
got out,but the sample was too sn~ll for statistical analysis.There
was a statistically significant increase in the strengths of reactions
when sudden loud noises were produced by the bus or visitors.The loud
noises may have been frightening because they were unexpected.Loud
noises also have an evolutionary importance to sheep as a danger signal
becaus e of th e preva 1ence of fa 11 i ng roc ks,rocks 1ides,and a va 1anc hes
in sheep habitat.Dixon (1938),Geist (1971c),and Jones et at.(1963)
all noted that wild sheep become alert to the sounds of falling rocks
and snow,but respond by moving only if in danger of being hit.Welles
and Welles (1961)found that bighorns in Ueath Valley learned to ignore
horns,gunshots,shouts,motors,and car noises.
The nature of the road-related human activities may facilitate the
habituation of the sheep to people on foot.People getting out of
buses and standing on the road are generally in full view of the sheep
and usually do not have enough time before the bus leaves to approach
the sheep close enough to frighten them.~lany investigators have found
sheep least disturbed by a man in full view below them and moving slow-
ly wtthout approaching directly (Dixon 1938,Jones et at.1963,Welles
and Welles 1950).Wolves are generally'successful in capturing sheep
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only if they surprise the sheep from above,at close range,and away
from escape terrain (A.Murie 1944).
The literature reveals that ewes with lambs are more sensitive to
disturbances than other classes of sheep (Jones et aZ.1963,A.Nurie
1944,Smith 1954).The experiences and temperament of the leader of a
group may also affect responses (Welles and Welles 1961).
129
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Brown Bear (Ursus aratos)
Population History,Numbers,Composition,Distribution
The br'o\,m bears of Mount McKi nl ey National Park have attracted the
attention of several naturalists and biologists in past years.Major
contributions to the knowledge of this poptilation appear in Dean (in
press),Dixon (1938),-A.Murie (1944,1961),and Sheldon (1930).The
bears of this region generally hibernate from October to April and the
young are born in the dens during mid-winter (A.Murie 1944,1961).
The breeding season extends from mid-May through mid-July (Dean,in
press,A.Murie 1961).
Dean (in press),using data from his earlier $tudies and from my
study,calculated minimum density estimates for the eastern half of
-2theparkof0.026,0.033,0.041,and 0.038 brown bears per km ,for
the years 1957, 1958,1959,and 1973,respectively.Although Sheldon
(1930)and A.Hurie (1944)did not make careful population estimates
for brown bears,their records on the frequency of sighting these bears
suggest that the population density probably was not greater than at
present.Thus,the numbers of brown bears in the eastern half of the
park in 1973-1974 probably equaled or exceeded the numbers present in
1906-1908 (Sheldon 1930),1939-41 (A.Murie 1944),and 1957-1959 (Dean,
in press).Pearson (1975),studying the Il nor thern interior grizzly
[brown bear]ecotype"at the K1uane Game Sanctuary in the Yukon,esti-
mated a population density of 0.037-0.043 brown bears per km 2 ,very
similar to the estimates for Mount McKinley National Park.Other esti-
mates of densities of interior brown bear populations from areas of
varying habitat quality include 0.045 bears per km 2 in Glacier National
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Park,Montana (Martinka 1974a),0.017 bears per km 2 in Yellowstone
National Park,Wyoming (Cm/an et al.1974),and 0.007-0.008 bears per
km 2 in the Brooks Range,Alaska (Curatolo and Hoore 1975,Quimby 1974).
Brown bears were commonly sighted from the shuttle-tour buses
during my study (Table 33).An average of 5.9 bears in 2.6 groups
were observed during each of the 70 round-trips.The average number
of "different"individuals Has 4.9.The maximum number of different
bears sighted on one trip was 14.Bears were sighted on 87%of all
trips.Matched-pair t-tests comparing numbers of bears observed from
the various types of buses,i.e.0400 tour,0700 shuttle,1500 shuttle,
on trips within two days of each other revealed no significant differ-
ences at P=0.05 in the numbers of bears observed from each type of bus.
The only seasonal trend in the numbers of bears sighted was a decrease
in numbers in August 1974 (Appendix A12).
Bears frequent,in varying concentrations,all areas along the
road.The Sable Pass area,between Tattler Creek and East Fork River,
produced the greatest concentration of sightings.Of all shuttle-tour
bear sightings,5%occurred between Riley Campground (Km 0.2)and
Tattler Creek (Km 60),34%between Tattler Creek and the East Fork
River (Km 70),and 61%between the East Fork River and Eielson Visitor
Center (Kin lO7).Table 34 gives an indication of the seasonal varia-
tions in use of different areas along the road.These variations are
probably related to the availability of various foods.A.Murie (1944)
outl ines four major phases in the food habits of the park I s bears,sug-
gesting that the bears feed on roots in the spring and early June,green
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Table 33.Number of brown bears observed during shuttle-tour trips,
Mount McKinley National Park,Alaska.
YEAR TOTAL NUMBER OF AVERAGE NUMBER OF BEARS AVERAGE NUMBER OF
,BEARS OBSERVED OBSERVED EACH TRIP "DIFFERENT"BEARS
OBSERVED EACH TRIP
Animals Groups Animals Groups Animals Groups
1973 279 122 6.8 3.0 5.7 2.5
1974 136 58 4.7 2.0 3.6 1.6
TOTAL 415 180 5.9 2.6 4.9 2.1
132
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Table 34.Distribution of brown bear observations during shuttle-tour trips by season and distance along road.Mount
rlcKinley National Park.Alaska.
AREA YEAR
LATE "lAY-JU~IE
SEASON
JULY AUGUST-EARLY SEPTEMBER
%of Trips Averaqe %of Trips Average %of Trips Average
Bears Observed Groups/Trip Bp.ars Ohserved Groups/Trip Bears Observed Groups/Trip
Riley Campground (km 0)1973 14 0.14 0 0 7 0.07
-Tattler Creek (km 60)1974 9 0.27 0 0 17 0.17
Tattler Creek (km 60)1973 33 0.80 83 1.83 29 0.28
-East Fork River (km 70)1974 55 1.00 83 2.00 . 8 0.08
East Fork River (km 70)1973 67 1.33 25 0.33 64 1.14
-kr;88 1974 54 0.64 17 0.33 17 ·0.17
km 88 1973 50 1.07 42 0.92 57 1.00
-Eielson (km 107)1974 36 0.73 50 0.83 25 0.42
TOT AL rlU~·lB ER TR I PS 1973 15 12 .14
1974 11 6 12
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134
vegetation,especially Equisetum and grasses,in June and July,berries
in late July through October,and roots again in late fall.All the
shuttle-tour bear sightings between Ril~y Campground and Tattler Creek
occurred in 1ate f\1ay and August.Bears were observed on Sable Pass,
between Tattler Creek and East Fork River,all through June,but heavy
use of this area began in late June,peaked in July,then decreased to
a low level in August.Since green vegetation,especially grass,is
the main food source for bears on Sable Pass,this use pattern conforms
with A.Murie1s (1944)data on the seasonal use of green vegetation.
Bear sightings between the East Fork River and Km 88 occurred chiefly
in June,with considerably fewer sightings in July,and an increase
again in August.Much river bar is visible from the road in this area
and during June brown bears often feed on the roots of Hedysarum grow-
ing on bars and flood plains CA.Murie 1944).Berries may also attract
bears to this area in August.Si~htings on the alpine tundra from Km
88 to Eielson were more variable,showing no distinct seasonal trends.
Of all the brown bears observed during the shuttle-tour trips,43%
occurred in dwarf shrub habitat,24%in alpine tundra,18%on gravel
bars and moraines,7%in tall willow and alder,3%in tussock-heath
wet tundra,2%in mixed talus and alpine tundra,1%on talus slopes,
and 1%in open spruce.
Data on the sex and age distribution of the bears observed during
the shuttle-tour trips appear in Table 35.These data include repeated
observations of certain individuals,as discussed below.Females with
young accounted for 64%of the groups sighted.Of all animals observed
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Table 35.Sex and age distributions of brown bears observed during
shuttle-tour trips,Mount McKinley National Park,Alaska.
SEX-AGE NUMBER OF BEAR OBSERVATIONS
1973 1974 TOTAL
Animals Groups Animals Groups Animals Groups
#of Singles 42 42 19 19 61 61
#of Pairs (Mating?)8 4 0 0 8 4
#of Females w/young 76 76 39 39 115 115
#of Cubs 57 28 3 2 60 30
#of Yearlings 90 45 25 12 115 57
#of Two-Year Olds 6 3 50 25 56 28
135
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28%were females \'lith young,56~~were young bears accompanying a fe-
male,15%were single bears,and 2%were in pairs,probably mating
associations.An attempt was made to identify individual bears ob-
served in Mount McKinley National Park,both from the road and in the
back"country,during the years 1957-1959 and 1973 (Oean,in press).The
methods are described .in Dean (in press)..This effort resulted in esti-
mates for 1957-1959 of a population composition of 22%females with
young,39%young accompanying females,and 39%single bears.For 1973
these figures were 225%,45%,and 30%,respectively.Thus,the single
bears seem to be under-represented in the shuttle-tour observations.
Excluding the young bears following a female,a minimum·of 71%of
the s'ightings were of females.This figure includes only females with
young and the sightings of one single bear known to be female.F.C.
Craighead and J.J.Craighead (1966)reported sex ratios for tagged
brown bears in Yellowstone National Park favoring males 2:1 in cubs,
but favoring females in adult bears over 4 years old.Pearson (1975)
found a sex ratio for all his captured bears with home ranqes within
his Kluane Game Sanctuary study area that favored males 56:44,but he
does not discuss any relationship to age.
In all shuttle-tour observations of families,26%included spring
cubs,50%included yearlings and 24%included two-year olds.A skew
towards one age group of younq occurred each year.In 1973,37%of the
young observed were spring cubs,59%were yearlings and 4%were two-
year olds (n=153)while,in 1974,these figures were 5%,31%,and
64%,respectively (n=78).Dean's (in press)data from the park,in
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1957-1959,also showed a skew towards one age group of young each year.
The average number of young per fami ly for all shutt!e-tour obser-
vations was 2.0 (n=115).A total of 64 different families were iden-
tified in the park during the years 1957-1959 and 1973.The average
litter size for these four years combined was 1.81 for cubs,1.83 for
yearlings,and 1.67 for two-year olds (Dean,in press).Average litter
sizes for various brown bear populations reported in the literature
range from 1.60 to 2.36,with the coastal brown bears producing the
higher averages (Lentfer et aZ.1966,Martinka 1974a,Mundy and Flook
1973,Pearson 1975,Quimby 1974,Troyer and Hensel 1964).Group sizes
for all sightings of bears from the shuttle-tour buses appear in Table
36.Except for four observations of pairs which may have been mating
associations,all groups larger than one consisted of a female with
young.
An additional factor of importance is the long life potential of
the brown bear.Of all the brown bears legally killed in the game
management unit north of McKinley Park from 1970-1975,13%were 13.4
years old or over (Valkenburg 1976).Curatolo and Moore (1975)re~ort-
ed a mean age of 11.2 years for a population in the Brooks Range.
Several authors found wild brown bears that were 20-25 years of age
(Curatolo and Noore 1975,Mundy and Flook 1973,H.V.Reynolds 1974).
The significance of the age and sex data as related to possible human
disturbances will be discussed below.
In 1973,we placed some emphasis on observing bears and learned to
individually recognize several families and a few single bears.Family
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Table 36.Sizes of brown bear groups observed during
shuttle-tour trips,1973 and 1974,Mount
McKinley National Park,Alaska.
BEAR OBSERVATIONS
GROUP SIZE Animals Groups
Number (Percent)Number .(Percent)
1 61 (15)61 (34)
2 10 (2)5 (3)
3 336 (81)112 (62)
4 8 (2)2 (1)
TOTAL 415 180
138
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139
.
composition,color,size,facial characteristics,location,and any
distinctive physical features were used for identification purposes.
Single bears were much more difficult to identify than families.Table
37 gives the number of sightings and the range of dates of the sight-
ings of individually identified groups.These are minimum sighting
records,since many other observations could have been of these same
bears,but positive identification was not possible so the observations
were not included.These data indicate that a few families occupy home
ranges adjacent to,or including,the road,and that these families are
sighted many times throughout the summer.Seven individually recog-
-
nized famil ies accounted for a minimum of 75%of the 1973 sh.uttle-tour
observations of families,46%of all groups and 62%of all animals
sighted.Four of these families were observed on dates from June
through August.One identifiable single bear,observed from May
through August 1973,accounted for at least 28~~of the 1973 shuttle-
tour observations of single bears.Another 33%of the 1973 single
bear observations were made on Sable Pass,and I believe these obser-
vations involved chiefly two individuals.Thus,seven families and
three single bears accounted for at least 68%of the groups and 71%
of the animals observed,in 1973,on the shuttle-tour trips.At least
three of the families and one single bear were observed in both 1973
and 1974.Most observations of each given individual were confined to
a section of the road 15 km or less in length.A.Murie (1961)noted
that individual brown bears in the park possess definite,but overlap-
ping,home ranges,that may be used in successive years.He reports
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Table 37.Observations of individually recognized brown bears,Mount McKinley National Park,Alaska.
BEAR YEAR NUMBER OF NUMBER OF DAYS RANGE OF NUr~BER OF NUMBER OF TOTAL TOTAL NUMBER TOl~L RA~:GE
SHUTILE-TOUR OBSERVED ON DATES OTHER OTHER DAYS NUMBER OF DIFFERENT DAYS OF DATES
OBSERVATIONS SHUTILE-TOUR (MONTH/DAY)OBSERVATIONS OBSERVED OBSERVATIONS OBSERVED (MO~lTH/DAY)
FaMily #1 in 1973,1973 13 12 5/26-8/17 23 22 36 32 5/24-9/25
fe":'31 e I'li th blO
yearlings 1974 4 3 6/28-8/31 5 5 9 8 6/28-8/31
Fa~i1y =2 in 1973,1973 9 8 6/24-8/24 12 12 21 17 6/24-9/1
fe~,a 1e ~/;th b/o
cubs
Fawi1y #3 in 1973,1973 2 2 6/29-7/1 6 6 8 8 5/30-8/30
fe~ale with three
cubs 1974 6/14 6 6 7 6 5/28-8/22
Fa~i1y =4 in 1973,1973 15 10 5/27-8/18 12 10 27 18 5/22-8/1e
fera 1e <Ii th two
yearl i ngs
FaMily =5 in 1973.1973 4 2 8/24-8/31 7 7 11 8 8/24-9/23
fe:7a 1e wi th two
cubs 1974 0 0 10 10 10 10 5/26-9/2
,Far:i1y =6 in 1973,1973 14 11 6/23-8/25 4 4 18 14 6/23-8/25
fe:"a 1e I'li th two
year1 i ngs
Farily =.7 in 1973,1973 0 0 3 3 3 3 8/7-9/9
fe~a1e with three
cubs
FaT,ily =8 in 1974,
fe:::a 1e wi th two
cubs 1974 .2 2 7/18-7/31 3 3 5 5 6/20-7/31
Single #1 1973 12 8 6/12-8/17 9 9 21 17 5/29-9/3
1974 3 2 6/8-6/14 3 3 6 5 6/8-8/27
Sable Pass Single 1973 14 9 6/3-7/10
Bears (~ost obser-
vations of t~/O
bears)1974 8 6 5/26-7/23
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141
that a sow with cubs often 1imits its movements to an area 1ess than
19 km in diameter.Four families observed frequently by Dean (in
press)in the park in 1957 possessed minimum home ranges of 15-26 km 2•
Seasonal home ranges are sometimes much smaller.Radio-tracking stu-
dies conducted on brown bears in other areas have shown that single
males use larger home ranges than females (Curatolo and Noore 1975,
Pearson 1975),that individuals may use the same home range in succes-
sive years (F.C.Craighead and J.J.Craighead 1972a,Curatolo and
Moore 1975),and that long distance movements are sometimes made,
especially by bears homing after a transplant attempt (Steve Buskirk,
pers.comm.1974,F.C.Craighead and J.J.Craighead 1972a,Mundy and
Flook 1973).
Human Disturbance
General Avoidance
Brown bears were observed in all the study plots,illustrating
that this species finds sui~able habitat over much of the park.Mini-
mum density estimates for the plots range from 0.01 to 0.41 bears per
km 2 (Table 38).These estimates are generally higher than the minimum
density estinmtes of 0.026-0.041 bears per km 2 for a much more exten-
sive section of the eastern part of the park made by Dean (in press)
for the years 1957-1959 and 1973.This difference may result from the
greater difficulties of identifying all the bears in a larger area and
from the fact that the larger area may include more unsuitable habitat.
Dean's study,like mine,concentrated on the area near'the park road,
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Table 38.Average number of brown bears observed in study plots,Mount t>1cKinley National Park,Alaska.
PLOT AVERAGE DAILY AVERAGE DAILY NO.OF
TOTAL OBSERVATIONS FIRST SEARCH OBSERVATIONS Dft,YS
AN IriAlS
Actua 1 Densi t~Actual Dp.f1S i t 2 OBSERVED
Number (per km )Number (oer km )OUT OF
FIVE
Animals Groups Animals Groups Animals Groups Animals Groups
Igloo Road 1973 0.40 0.40 0.03 0.03 0.40 0.40 0.03 0.03 2
Igloo Road 1974 0.80 0.20 0.05 0.01 0 0 0 0
Igloo Off-Road 1973.2.20 1.40 0.12 0.07 1.20 1.00 0.06 0.05 4
Igloo Off-Road 1974 2.20 1.20 0.12 0.06 2.20 1.20 0.12 0.06 3.
Hi ghway Road 1973 0.20 0.20 0.01 0.01 0.20 0.20 0.01 0.01
Hi gh~lay Off-Road 1973 0.60 0.60 0.05 0.05 0.60 0.60 0.05 0.05 3
Stony Road 1973 0.80 0.40 0.06 0.03 0.80 0.40 0.06 0.03 2
Stony Off-Road 1973 1.20 0.40 0.12 0.04 1.20 0.40 0.12 0.04
Sable Road 1973 8.40 3.60 0.41 0.17 5.00 1.40 0.24 0.07 5
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and comparisons of several population parameters already discussed
earlier reveal no dramatic changes in the population between the years
1957-1959 and 1973,despite the great increase in visitation and traf-
fic level s.
The data from the paired plots (Table 38)show consistently
higher bear densities in the off-road plots compared to the road plots.
A n~tched-pair t-test comparing all first search densities of bears for
all observation periods in the backcountry plots with the respective
densities for the paired road plots (Appendix A8)shows significantly
more bears were observed in the backcountry plots (t=10.7,P<0.005).
It seems that some bears may be avoiding the vicinity of the road,
despite the fact that many bears are sighted from the road.Some sin-
gle bears,in particular,may be avoiding the road.As already men-
tioned,single bears appeared to be under-represented in the bus
sightings,compared to their representation in the population in gen-
eral.I observed single bears in the backcountry more frequently than
near the road.Excluding one habituated individual,single bears were
sighted within 100 m of the road on only three occasions during the
shuttle-tour trips.
However,the highest density of bears occurred in the unpaired
Sable Road Plot,an area of unique,high quality bear habitat and a
special managment unit where visitors are not permitted to move off
the road.The special restrictions were initiated in the early 1950's
because park personnel felt that over-persistent photographers were
causing bears ·to move away from the vicinity of the road.In this
[
plot now,with the restrictions,bears even within 200 m of the road
seldom reacted visibly to human activities on the road.However,in
144
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the nearby Igloo Road Plot,\"here much human activity occurs off the
road,two of four groups of bears sighted during plot observation per-
iods moved around a mountain and out of view of the road in response
to human activities.One group was responding to hikers and the other
to a passing shuttle bus.
Reaction Distances
The distribution of bears sighted during the shuttle-tour trips
in relation to distances from the road appear in Table 39.Numbers
observed generally decreased with distance from the road,and thus,
with visibility.Individual bears varied in their tolerance to proxi-
mity to the road.Ten of the thirteen observations of single bears
within 100 m of the road are accounted for by one recognizable indi-
vidual,which frequently fed on vegetation within a few meters of the
road and showed no concern as buses and cars passed and stopped.
Table 39 and Figure 13 show the reaction patterns of bears at
varying distances from the road.A G-test (Sokal and Rohlf 1969)run
on the reaction patterns of bears at 100 m intervals from the road up
to 500 m shows that there is a significant (G=75.7,P<O.005)change in
the reaction pattern that is related to distance from the road.For
all the shuttle-tour bear observations for which reactions could be
classified,78%of the animals exhibited no visible reaction to the
presence of the bus and people,7%gave mild reactions,and 15%gave
r tl~
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146
751-1000
N=33N=25
-N=61 -~
N=34-N=37 ,-
N=65 r-
Reaction
-0 None-IZJ Mild
II Strong
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20
(/)
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o
0-100 101-200 201-300 ~1-400 401-500 501-750
Distance from Rood (m)
100
~40
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Figure 13.Reactions of brown bears,observed Rt va~ying distances
from the road during shuttle-tour trips,to the bus and
visitors,1973-1974,Mount tlcKinley National Park,Alaska.
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14-7
strong reactions.For brown bears within 200 m of the road,these
percentages "'Jere 56%,12%,and 32%,respecti ve ly.Ei ghty-th ree percent
of the bears that reacted to the bus and visitors were within 200 m of
the road,and 96%were within 400 m.
Very little information is available in the literature on the
flight distances of brovm bears.Both Klein (1974)and t1cCourt et aZ.
(1974)found brown bears more sensitive to aircraft overflights than
caribou or moose,although the reactions of the bears were quite
variable.rkCourt et aZ.(1974)reported that 8 of 10 bears subjected
to ovel~flights above 1000 ft (300 m)still exhibited "significant"reac-
tions to the aircraft.Quimby (1974)determined that 32%of the bears
sighted within 1 mi (1.6 km)of his aircraft were already running away
when sighted.~1ost of the bears \-lith which Quimby was working had
already been subjected to drugging and tagging operations utilizing
ai rcraft.
The distance at which a grizzly bear may detect and react to a man
on foot is poorly documented.F.C.Craighead and J.J.Craighead
(1966),after years of experience around brown bears,felt that the
bears often could not discern the form of a motionless man beyond
100 yd(90 m).Pearson (1975)concluded that brown bears usually ex-
hibit little alertness and are easily approached closely by a man,if
no scent reaches them.In areas where human hunting is prohibited
adult brown bears have few enemies excent other bears and constant
alertness may not be necessary.
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148
Disturbance Frequencies
The frequency of road-related disturbances to bears may be calcu-
lated by combining the frequency data for vehicles passing through the
Igloo Canyon-Sable Pass section of the road (Fig.5)with the reaction
pattern of bears within 200 m of the road (Table 39).This calculation
predicts that an "average"bear within 200 m of the road exhibits 0.8
mild responses and 2.2 strong responses to vehicles and visitors each
hour from 0600-2100 hours.Since no intensive observations were made
on bears,the exact disturbance frequencies for individual bears cannot
be given.The calculation above is meaningful only if a bear remains
within 200 m of the road for an extended period.This situation may
actually occur quite often.Bears were frequently resighted during
the shuttle-tour trips,indicating that they commonly remain in view of
the road for hours at a time.Forty-five percent of the bears sighted
from the bus while the bus was traveling east were resighted on the re-
turn trip west the same day.Fifty-six percent of the resighted bears
were within 200 m of the road during the first sighting.Fifty-three
percent of the resightings occurred 2-5 hr after the first sighting;
the rest occurred within 2 hr of the first sighting.
No data are available on the usual frequency of activity interrup-
tions caused by non-human-disturbances.The fact that the brown bear
is a top level carnivore and its lack of alertness suggest that non-
human disturbances are probably infrequent.Females with young were
occasionally observed running from other bears.
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Relationship of Reaction to Type of Disturbance
Table 40 and Figure 14 present the data from the shuttle-tour
trips on the actions of the buses and visitors and the responses of
brown bears within 200 m of the road to various types of human activi-
ties.The buses stopped for almost all bears sighted,passing only 5%
of the animals.The buses stopped,with the people remaining on the
bus,for 70%of the bears and stopped,with people getting out of the
bus,for 25%of the bears.These figures for bears within 200 m of
the road were 2%,66%,and 32%,respectively.People were not observed
to move off the road towards bears during these trips,although photo-
graphers were seen to do so at other times.The people accidently or
purposely produced particularly loud noises while observing 10%of all
animals,and while observing 16%of the animals within 200 m of the
road.A comparison of shuttles and tours shows that visitors on the
shuttles got out to observe 33%of the bears sighted,but visitors on
the tours got out to observe only 8%of the bears sighted.
Chi-squared tests were used to compare the reactions bears within
200 m of the road exhibited towards the various actions of the bus and
visitors.No significant relationships at the P=0.05 level were found.
The response patterns (none:mild:strong)varied little,regardless of
whether the bus simply stopped with the people remaining on the bus or
if people got out of the bus and stood on the road.Loud noises caused
an increase in the perentage of animals reacting strongly,but sample
sizes were small.A.Murie (1961)commented that producing loud noises
is a good way to drive away brown bears.It is possible that bears more
(J
Table 40.Reactions of brown bears,observed within 200 m of the road during shuttle-tour trips,to various actions of the bus and
visitors;percent of animals ~nd groups,1973-1974,Mount McKinley National Park,Alaska.
ACTION OF BUS KllOWN REACTIONS OF GRIZZLY BEARS TOTAL TOTAL OBSERVATIO~S
AJ'lD VISITORS KNOHN KNmJN AN 0 UN KNmm
NONE MILD STRONG REACTIONS REACTIONS
~;Grand %Grand
Percent Percent Percent Percent Percent Percent Number Number Number Number Total iotal
Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups
Pass >-15 mph 100 100 0 0 0 0 4 2 2 3
Pass <15 mph 0 0 0 a 0 0 0 a a 0 0 0
Stop,peoole 59 54 12 18 29 28 90 39 100 43 54 56
re;r:ain on
bus,quiet
Stop,people 45 38 10 25 45 38 20 8 21 9 11 12
remain on
bus,noisy
Stop,people,64 67 12 19 24 19 50 21 50 20 27 26
off bus on
road,quiet
Stop,peoole 50 50 0 0 50 50 6 2 9 3 5 4
off bus on
road,noisy
Stop,people 0 0 0 0 0 0 0 a 0 0 0 0
wal;:off
road ,.qui et
Stop,people 0 0 0 a a 0 a a a a 0 a
~:a 1k off
road,noisy
TOTAL 59 56 11 18 30 27 167 .71 184 77
Note:"Groups"i.lay total over 100%,since some qroups ~Iere counted under more than one reaction class when various animals in the
group reacted differently.Raw numbers for animals aQd'qroups are presented in Appendix Table A28.
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Visitors Remain On Bus
(Quiet)(Noisy)
Bus Stops
151
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Figure 14.Reactions of brown bears,observed within 200 m of the
road during shuttle-tour trips,to various actions of
the bus and visitors,1973-1974,Mount McKinley National
Park,Alaska.
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152
than 100-200 m from the road do not detect the details of the activities
of buses and visitors.The brown bear's eyes are relatively small and
their eyesight is apparently poor,although very little discerning in-
formation is available on the senses of bears.Russell (1967)felt
that bears can see little beyond 300 yd (275 m),and F.C.Craighead
and J.J.Craighead (1'966)felt that brown bears cannot identify a
.motionless man by sight beyond 100 yd (90 m).However,t1acPherson
(1965)proposed that vision in bears is similar to manis,but with
less ability to perceive movements at a distance.Sight seems to play
a supporting role to the other senses and the sighting of objects or
movements,alone,does not stimulate flight in bears (Johnson 1972,
Pearson 1975).The ears of brown bears are small and quite rigid,un-
like the large,mobile ears of most ungulates and many carnivores.F.
C.Craighead and J.J.Craighead (1966)and Krott and Krott (1963)con-
cluded that hearing in brown bears is keen and important,although
MacPherson (1965)and Johnson (1972)felt it plays only a secondary
role.There are few published comments on distances at which bears
can detect sounds.Linderman (1974)noted a brown bear responding,
apparently to the sounds of an aircraft,before Linderman himself
could hear it.He also noted that bears,especially those that had
been tagged previously,sometimes began fleeing when aircraft could be
heard but not seen.Pearson (1975)also observed this type of learned
response to aircraft sounds.Most authors agree that the bear's sense
of smell is acute and the most important sense in warning it of danger
(F.C.Craighead and J. J.Craighead 1966,Dixon 1938,Johnson 1972,
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153
Kratt and Kratt 1963,MacPherson 1965).Johnson (1972)often observed
brown bears scenting a man at distances of 150 yd (130 m).Human smell
alone may cause precipitous flight in bears.F.C.Craighead and J. J.
Craighead (1972a,1972b)related an instance where tracks in snow re-
vealed that a grizzly coming upon a human trail had urinated,and then
had run for at least 200 yd (180 m).During mv study,I watched from
a ridgetop as a grazing brown bear came upon my trail in the valley.
The bear responded to the scent by running at least 4 km across a valley,
through a stream,over a steep talus ridge,and out of view.Hasselborg
(reported in Schoonmaker 1968)observed a bear responding to human scent
on a trail 14 hours after the man had passed.
Brown bears seem to learn easily and modify their responses to sim-
ilar stimuli in varying situations.J.J.Craighead and F.C.Craighead
(1972)found that their tagged bears apparently ignored human scent
while feeding in garbage dumps but that the same individuals would flee
after receiving a human scent in an area away from the dumps.I noted
a similar dichotomy in Mount McKinley National Park.Certain families
I had learned to recognize and had observed showing apparent habitua-
tion towards people and traffic on the road fled from me when I met
them in the backcountry.
Relationship of Reaction to Sex and Age
Separate analyses of the distribution in relation to distance from
road and the reaction patterns for single bears,females with cubs,and
females with yearlings or two-year olds revealed only one difference
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154
between these groups;single bears exhibited fewer strong reactions
(Table 41).This result may be misleading.As mentioned earlier,10
of the 13 observations of single bears within 100 m of the road were
all of one individual.Thus,single bears other than this individual
were seldom observed within 100 m of the road.
Many authors (Herrero 1970,Mundy and Flook 1973,Riegelhuth 1966)
have concluded that female bro\tln bears with young are more likely to
attack during disturbing situations than single bears.This behavior
may change with the age of the young (Pearson 1975).However,attack
behavior was seldom observed by us in road-related visitor-bear inter-
actions and this difference was not apparent.Barnes and Bray (1967)
noted an interesting sex-related difference in black bears in Yellow-
stone National Park;the campgrounds were frequented mostly by adult
males and the roadsides mostly by females and yearlings.
Relationship of Reaction to Behavior Before Disturbance
The behavior before disturbance,when known,for bears sighted on
the shuttle-tour buses appears in Table 42.The apparent behavior~l
differences between bears within 200 m of the road and those further
away probably results simply from visibility problems.Traveling bears
are easier to sight at a distance than bears engaging in other activi-
ties,while lying bears are particularly difficult to sight at a dis-
tance.
Table 43 presents the data on the relationship between reaction
and behavior before disturbance for bears within 200 m of the road.
r"rJ IT ',r ,c:r~('-r"]c-:-n c:-J CJ CTJ r J rr----'CJ ::-1 l'J rJ :-J ,:-l :-l r-J)L1 u.,..,:'".J
Table 41.Reactions of singl e brown bears and famil ies,observed during the shuttl e-tour trips,
to the bus an~visitors,1973-1974,~lount fvlcKinley National Park,Alaska.
DISTANCE REACTION OF BROWN BEARS TO BUS AND VISITORS TOTAL
FRON ROAD KNmm
(01)NONE NILU STRONG REACTIONS
Percent Percent Percent Percent Percent Percent Number Number
Animals Groups Animals Groups Animals Groups Ariimals Groups
SINGLE BEARS
0-100 46 46 54 54 0 0
13 13
101-200 62 62 12 12 25 25 8 8
201-400 100 00 0 0 0 0 17 17
>400 100 00 0 0 0 0 19 19
Total 57 57
FAmLIES
0-100 49 47 10 17 42 43 88 30
101-200 79 79 5 5 16 16 57 19
201-400 86 85 10 12 4 4 79 26
>400 97 96 0 0 3 4 87 29
Total 311 104
Note:"Groups"may total over 100%,since some groups were counted under more than one
reaction cla s when various animals in the group reacted differently.
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Table 42.Behavior before disturbance of brown bears observed during shuttle-tour trips,1973-1974,Hount McKinley National Park,Alaska.
OISTAHCE
FRO:i ROAD
(m)FEED
BEIiAVIOR OF BROWa lHARS GHORl OISTURBAliCl
FEED-WALKa LIE ST';IW TRAVEL b PLAY HU!tT
TOTAL
li~X::'lR
N.IIJ"'~S
Number (Percent UUr:'ber (Percent Number (Percent Number (Percent liun,ber (Percent rlumber (Percent !lumber (Percer.t
Anir.1clls Anir.lills)Animals Animal s)Aninlills Animals)Animal s Anir.lills)Animals Animals)Animals Animals)Animals Anirr.als)
0-200 76 (52)24 (16)22 (15)0 (0)13 (9)8 (6)2 (1)](~
>2JO 95 (40)24 (12)11 (6)3 (2)53 (27)4 (2)7 (4)197
TOTAL 171 (50)48 (14)33 (10)3 (1)66 (19)12 (4)9 (3)342
a Feeding while traveling at a walk.
b Movement at a walk,trot,or run.
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Table 43.Reactions of brown bears,observed within 200 m of the road during shuttle-tour
trips,to the bus and visitors as related to behavior before the disturbance,
1973-1974,Mount McKinley National Park,Alaska.
BEHAVIOR REACTION OF BROvJN BEARS TO BUS AND VEHICLES TOTAL
BEFORE NU~IBER
DISTURBANCE NONE NILD STRONG ANINALS
Number (Percent Number (Percent Number (Percent
Animals Animals)Ani/llals Animals)Animals Animals)
FEED OR HUNT 55 (71 )8 (10)15 (19 )78
FEED-HALKa 20 (83)3 (12)1 (4)24
LIE 17 (81)4 (19 )0 (O)21
TRAVEL b 4 (40)0 (0)6 (60)10
PLAY 8 (100)0 (O)0 (0)8
TOTAL 104 15 22 141
a Feeding while traveling at a ~~lk.
b Movement at a walk,trot,or run.
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This table includes only animals for which the behavior before distur-
bance was determined.Sample sizes are insufficient for statistical
analysis,so the data are only useful as a preliminary indicator.Ly-
ing bears seem less responsive than feeding or hunting animals,while
traveling individuals appear most sensitive.Possibly,as with the
caribou,some of the traveling bears were still responding to previous
disturbances and were particularly sensitive for this reason.
Behavior Patterns Related to Disturbance
The maximum behavioral reactions of brown bears in response to
buses and visitors appear in Table 44.Of those bears sighted within
200 m of the road 58%gave no visible reaction and did not interrupt
their normal activities in response to the human activities.This fig-
ure indicates a high degree of habituation in the individual bears in-
volved.Bears within 200 m of the road were frequently observed engag-
ing in such activities as nursing,playing,and sleeping,which suggest
security and relaxation.Only 8%of the anin~ls within 200 m of the
road lifted or turned their heads and looked at the human activities on
the road without subsequently moving away;10%moved away at a walk and
23%at a run.Such movements were seldom over 100 m,but occasionally
bears moved 200 m or more and out of view of the road.These short
movements differ from my experi ences with bears in the backcountry,
where bears that had become aware of my presence usually ran out of
view and sometimes ran 2 km or more.Bears fleeing from aircraft have
often been observed to hide,when patches of dense vegetation were
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Table 44.Maximum behavioral reactions of brown bears,observed during shuttle-tour trips,to buses
and visitors,1973-1974,Mount McKinley National Park,Alaska.
DISTANCE ~1AXI~IUM REACTION OF BROWN BEARS TO BUS AND VISITORS TOTAL
FRO~I ROAD NUivlBER
(m)NONE WATCH ~JALK TROT-RUN ANIHALS
Number (Percent Number (Percent Number (Percent Number (Percent
Animals Animals)Animals Animals)Animals Animals)Animals Animals)
0-200 98 (58)14 (8)(17)(10)38 (23)167
>200 195 (93)5 (2)0 (0)9 (4)209
TOTAL 293 19 17 47 376
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available (Erickson 1965 t Klein 1974 t Pearson 1975 t Quimby 1974).
The Itypica1"response of a brown bear that has detected a strange
or alarming object is to stand upright on its hind legs t sway its head t
inhale deeplYt and perhaps circlet to try to get a scent (Johnson 1972 t
Pearson 1975).If a human scent is detected the animal often retreats
hastily (A.Murie 1961 t Pearson 1975).These scent-detection behavior
patterns were seldom observed in response to human activities on the
road in the park.We observed bears standing upright on only five oc-
casions;once by a female with young trying to cross the road in the
midst of several vehicles t once by a female with young watching a
camper-truck containing a loudly barking dog t once by a yearling when
a bus driver honked a horn t and once each by a cub and a yearling just
watching toward vehicles on the road •
Attacks on humans by brown bears are uncommon.Herrero (1970)
surnnarized records for all North American national parks containing
brown bears and found an injury rate from brown bears of five persons
per year or one person per 1.5 million visitors.He reported only five
brown bear-caused deaths in the parks in 97 years.In the 14 years
from August 1961-August 1975 in Nount McKinley National Park,six
attacks occurred,none of them fatal (Buskirk 1975 t Cauble 1975).At
least two of these attacks were provoked by the close approach of
photographers,and at least five of them involved females with young.
After an intensive literature review Riege1huth (1966)concluded that
the brown bear's temperament is chiefly defensive t not aggressive,and
that attacks generally occur under one of three circumstances:1)a
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close approach to a female with young,2)a close approach to a bear
at a favored food supply,or 3)surprising a bear at close range.Even
under these'circumstances a bear usually runs.Herrero (1972)attri-
butes the defensiveness of brown bear females with young to the fact
that young rarely c1 imb trees,are often in open country,and depend on
the female for protection.Feeding on garbage or human foods may also
make a bear dangerous (Johnson 1972,Nartinka 1974b).The defensive
behavior of bears seems similar in bear-bear and bear-man interactions
(Barnes and Bray 1967).Bluff charges are much more common than actual
attacks (Barnes and Bray 1967,F.C.Craighead and J. J.Craighead
1972a,Russell 1967).No attacks were recorded during interactions of
bears and vehicles or persons near vehicles along the park road during
my study,even though vehicles and visitors were often within 50-100 m
of females with young.Apparently these females did not consider the
situation threatening,since the people did not move off the road to-
ward them.Also,the bears have not become accustomed to receiving
food from people,a situation which often leads to injuries.On one
occasion I witnessed a short bluff charge by a two-year-old towards a
person standing by the open door of a car.
We frequently observed bears crossing the road,even when vehicles
were nearby.They occasionally traveled along the road.I observed
one family travel at least 2 km on the road,once making a detour around
several parked cars and then returning to the road.
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Influence of Roadside Vegetation
As described previously,for most of its length the road influences
both the species composition and phenology of a narrow strip of vegeta-
tion on either side.This roadside vegetation includes several species
of grasses,herbs,and Equisetum which are utilized as food by brown
bears.On 11 occasions I observed bears feeding on this narrow band of
vegetation,apparently drawn to the road or remainin,g by the road be-
cause of its presence.All such observations occurred between June 10
and June 20 each year.From gross examination it appeared that the
roadside vegetation began grO\'/ing and IIgreening-upll somewhat earl ier
than the other vegetati on,probably because of snow removal,quickened
melt because of dust,and drainage patterns related to the road.
Apparently,the bears at times take advantage of this early growth.
Three different families and one single bear were involved in these
observations.The one single bear,a particularly habituated individ-
ual,accounted for seven of these observations and,unlike the other
bears,she frequently continued feeding on the roadside vegetation as
vehicles stopped or passed.
Influence of Garbage
Nount NcKinley National Park has only one garbage dump presently
in use within its borders.In conjunction with this study,Chip Oown-
ing monitored \'/ildlife activity at this dump and has reported the re-
sults in a manuscript to NPS (Downing 19.75).During the summer of
1974,brown bears frequented the dump throughout the summer;17
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different individuals were identified and a few more probably used the
area.Garbage feeding adversely affects the distribution and activities
of a wild population of bears and is a lamentable situation.The prob-
lem is still small in Mount tkKinley National Park compared to such
areas as Yellowstone National Park where visitation levels are much
higher (Cole 1972,J. J.Craighead and F.C.Craighead 1972b,Martinka
1974b).Since my study was completed,NPS has already implemented
plans to eliminate garbage,in the dump and at the campgrounds,as a
source of food for bears (Steve Buskirk,pers.comm.1976).
Summary and General Discussion
The minimum density of brown bears in the eastern half of the park
in 1973 was estimated at 0.038 bears per km 2 (Dean,in press).The
population lev~l appeared to be as high or higher in 1973 as in past
years when visitor numbers were much lower (Dean,in press,A.Murie
1944,Sheldon 1930).Bears were sighted on 87%of the shuttle-tour
trips.The sex and age data for the bears observed from the buses
indicate:1)a predominance of family groups,2)a predominance of
females for those bears sighted that are over two years of age,and
,3)a skew towards one age class of young each year.A few individual
families and single bears were sighted repeatedly from the road.Seven
families and three single bears accounted for a minimum of 68%of the
groups and 71%of the animals observed during the 1973 shuttle-tour
trips.Individuals may live up to 20-25 years (Curatolo and Moore
1975,t·1undy and Flook 1973,H.V.Reynolds 1974)and home ranges may be
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used in successive years (f.C.Craighead and J.J.Craighead 1972a,
Curatolo and Moore 1975).Thus,certain individuals are frequently in
contact with human activities along the road and may habituate or be-
come stressed,largely depending on the behavior of the humans.Impor-
tant factors in the biology of brown bears to consider when evaluating
the potential effects of disturbance include low population densities,
low reproductive potential,restricted home ranges,and long life.
The plot data and shuttle-tour data suggest that some bears,par-
ticularly single bears,may be avoiding the vicinity of the park road.
Single bears,particularly males,usually use larger home ranges than
families (Curatolo and Moore 1975,Pearson 1975).A single bear with
a larger home range may come in contact \'Jith the road less frequently
and have less opportunity to habituate than families with small home
ranges.A larger home range would also allow more areas for a bear to
use in preference to the vicinity of the road.Individual bears whose
home ranges extend outside the park,especially during hunting season,
may exhibit more wariness towards man under all circumstances.Past
experiences in the park,which led to the establishment of the Sable
Pass Wildlife Restricted area,also indicate that bears may avoid the
vicinity of the road under some·circumstances.faro and Eide (1974)
reported that increasing numbers of photographers at McNeil River falls,
Alaska,caused brown bears to modify the time and location of their
fishing activities to reduce contact with humans.They also felt that
some bears may have abandoned the falls completely.
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However,several families and a few single bears have become
habituated to the road and traffic in NcKinley Park,and these bears
were the ones most commonly sighted from the shuttle-tour buses.The
threshold distance \vhere 50~~of the animals exhibited strong responses
occurred between 50 and 75 m from the road.The threshold distance for
50%visible response occurred between 100 and 200 nl from the road
(Table 39).
The buses stopped for 98%of the bears sighted \'Jithin 200 m of the
road,while people got out onto the road to observe 32%of these bears.
During these trips people did not move off the road towards the bears •
No significant differences in the reactions of the bears were found
that were related to the type of activity of the bus and visitors.
There were large individual variations in responses,and these varia-
tions,coniliined with small sample sizes,may have obscured any general
patterns.Adult brown bears are usually not susceptible to predation
except by man and,occasionally,other bears.The danger from man to
North ~nerican bears was probably minimal until development of depend-
able rifles.Johnson (1972),after reviewing the accounts of explorers
in North America,concluded that bears had to learn to fear man and the
first explorers found them very unafraid and unwary.It is possible
that the brown bears must learn responses to various types of environ-
mental stimuli and possess few innate fear responses,as compared to
species that have evolved under the selective pressures of effective
predators.Perhaps,for this reason bears may show large individual
variations in responses to various types of stimuli,i.e.the
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components of approach,stopping,and visitors unloading from a bus.
Information from the literature discussed earlier also indicates that
the bear's senses may not detect the detail of visitors'activities
beyond 100-200 m (F.C.Craighead and J.J.Craighead 1966,Russell
1967)•
Bears occasionally seem to be attracted to the road to feed on the
roadside vegetation during June.During this period the roadside vege-
tation is often green and growing a1tho~gh the vegetation in the sur-
rounding area may not have begun to green.
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Red Fox (VuZpes vuZpes)
Population History,Numbers,Composition,Distribution
A.Murie (1944)made numerous general observations on the red fox
population in Mount McKinley National Park.He studied food habits,
movements,and den sites.He described this species as lI a bundant"
during the period of his studies.The only other study focusing on
the red fox in the park was conducted by Allison (1971)during 1968-
1969.She described den characteristics,activity patterns and
behavior.
The number of red faxes observed from the shuttle-tour buses dur-
ing my study appears in Table 45.The average number of animals ob-
served each trip was much lower in 1974 (0.4 animals/trip)than in
1973 (2.1 animals/trip).In 1974,faxes were observed on 34%of the
trips,while in 1973,they were observed on 63%of the trips.The
difference between the two years is accounted for by the large number
of sightings,74%of all animals,that occurred at or within 1 km of
known active den sites in 1973.Although there were active dens within
view of the road in 1974,they were much less well known by the bus
drivers.All 1974 sightings during the shuttle-tour trips were of
adults more than 1 km from kno\'tn den sites.The number of faxes using
the area may have actually decreased to a low level in 1974.The
number of active fox dens seemed to decrease,as discussed below.
During both summers,sightings were made more frequently during the
first half of the summer than during the second half.In 1973,the
average number of animals sighted each shuttle-tour trip before July 15
was 3.38,while after July 15 this number dropped to 0.80.For 1974,
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Table 45.Number of red fox observed during shuttle-tour trips,
Mount McKinley National Park,Alaska.
YEAR TOTAL NUI/lBER OF AVERAGE NUMBER OF
RED FOX OBSERVED RED FOX OBSERVED EACH TRIP
Animals Group Animals Group
1973 87 46 2.1 1.1
1974 13 13 0.4 0.4
TOTAL 100 59 1.4 "0.8
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these figures were 0.83 and 0.18,respectively.This decrease may be
related to decreased activity around den sites,movement of families
to new,unknown dens,and decreased daylight activity of adults,
associated with the increasing independence of the young.Foxes were
observed all along the park road and in all major types of habitat.
During my study,26 fox den sites were located within 1 km of the
road.The distances from the road and use of these dens during 1973-
1974 are shown in Table 46.In 1973,9 active dens were known within
400 m of the road.In 1974,5 active dens were known within 400 m of
the road.Some families used more than one known den site during a
summer;the number of different families known to use one or more dens
within 400 m of the road was nine in 1973 and four in 1974.A.Murie
(1944)reported that in 1940 he located 5 active dens along a 32 mi
(51 km)stretch of the park road,and knew the approximate location of
4 other dens in the same area.Allison (1971)was able to locate 20
dens known to be used sometime during 1967-1969,although she does not
report the distances between the dens and the road.
Pups,as well as adults,were frequently sighted from the road.
In 1973,59%of the fox sightings from the shuttle-tour buses were of
pups.In 1974,all the shuttle-tour sightings were adults,but I ob-
served many pups from the road in 1974 during mY intensive fox observa-
tions.The average litter size for eight families was 4.8 pups.
Ables (1975)reviewed the literature on home ranges and movements
of red foxes in North America.He concluded that the normal activities
of individuals and families in temperate habitats are generally
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Park road,1973-1974.
170
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DISTANCE NUMBER AND USE OF KNOWN RED FOX DENS TOTAL
FRO~1 ROAD KNOW~
(m)ACTIVE INACTIVE ACTIVITY UNKNOWN DENS
1973 1974 1973 1974 1973 1974
0-100 3 3 5 5 0 0 8
101-200 3 1 5 3 0 6
201-400 3 1 3 1 0 4
401-1000 2 4
2 3 4 1 8
TOTAL 11 9 7 16 8 1 26
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conducted in an area less than 8 km 2 and that individuals may use the
same home range or seasonal home range for several yearso A.Nurie
(1944)was able to recognize several individuals in McKinley Park,
each of which used a restricted home range over 2-4 yearso A.(·lurie
(1961)also noted foxes i~the park hunting 3-4 mi (4-6 km)from their
dens.Allison (1971)regularly observed adults hunting over a mile
(1.6 km)from their dens within the park.
Human Disturbance
General Avoidance
The plot data for foxes are presented in Table 47.A comparison,
using a matched-pair t-test,of all first search observation periods
in the backcountry plots v/ith the observations in the paired road plots
reveals a weak significant difference (t=1.7,P<O.05),with the density
of foxes in the road plots being higher (Appendix A10).Fourteen of
the 16 foxes observed during first searches were in the road plots.
These sightings were generally associated with active dens.In 1973,
the Igloo,Highway,and Sable Road Plots each contained a known active
den.No active dens were discovered in the off-road plots.Some in-
dividual foxes may be attracted to the road,as discussed below.There
is no evidence that a large percentage of foxes avoid the road as an
activity area or as a denning area,although individual differences do
occur.As discussed earlier,many active dens were in view of the
road,including one only 5 m from the road.Most pups are born before
~he park road is open to traffic in the spring,so no vehicular
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Table 47.Average number of red foxes observed in study plots,Mount McKinley National Park,Alaska.
PLOT AVERAGE DAILY AVERAGE DAILY i~UM3ER
TOTAL OBSERVATIONS FIRST SEARCH OBSERVATIONS OF UAYS
ANU1ALS
Number Density Number lJensity OBSERVED
(Per km 2)(Per km 2)OUT OF 5
POSSIBLE
Animals Groups Animals Groups Animals Groups Animal s Groups
Igloo Road 1973 1.00 0.60 0.06 0.04 1.00 .0.60 0.06 0.04 2
Igloo Road 1974 1.20 0.60 0.08 0.04 1.20 0.60 0.08 0.04 2
Igloo Off-Road 1973 0.20 0.20 0.01 0.01 0 0 0 0 0
Igloo Off-Road 1974 0 0 0 0 0 0 0 0
0
.Highway Road 1973 0.80 0.60 0.04 0.03 0.60 0.40 0.03 0.02 2
Hi gh\'/ay Off-Road 1973 0.20 0.20 0.02 0.02 0.20 0.20 0.02 0.02 1
Stony Road 1973 0 0 0 0 0 0 0 0 0
Stony Off-Road 1973 0.20 0.20 0.02 0.02 0.20 0.20 0.02 0.02 1
Sable Road 1973 0.20 0.20 0.01 0.01 0 0 0 0
'0
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disturbance occurs during the selection of whelping dens.However,no
abandonment of dens near the road was noted when the road opened or
when the volume of traffic increased.Allison (1971)noted that two
families moved from whelping dens to summer dens that were closer to
and more visible from the road.The red fox is well known for its
adaptability to environments modified by man (Ables 1975,Lloyd 1975,
Storm 1972).
Reaction Distances
The responses of foxes at various distances from the road to vehi-
cles and visitors are presented in Table 48.In addition to my general
definition of a "s trong"response given under "~~ethods,"if a fox en-
tered a den in response to a disturbance it was considered a strong
response.During my intensive observations of foxes at dens,animals
within 100 m of the road responded to vehicle and visitor activities
mildly 51%of the time and strongly 15%of the time.For animals
101-200 m from the road,these figures decreased to 27%and 1%,respec-
tively.However,the shuttle-tour data indicate greater responsiveness
in foxes within 100 m of the road but aw~y from den sites.These ani-
mals gave 45%mild responses and 36%strong responses.Probably more
of these animals were unhabituated than those observed at dens near the
road.Occasionally,foxes traveling or resting on the road showed no
visible response to vehicles or visitors also on the road,and they
often gave only mild responses.However,foxes 600 m from the road
were occasionally observed responding to passing vehicles.Few
~L 174
[
[Table 48.Reactions of red foxes at various distances from the-road to vehicles
and visitors,1973-1974,liount I.JcKinlcy Hational Park,Alaska.
r OISTArlCE REACTIONS OF RED FUXES TOTAL
i FROI-I ROAU iMHH::R
(m)riO/IE HILU STROHG AIHHALS
1 Number (Percent Ilur.lber (Percent r~umber (Percent
Animals Animals)An i,nd 1s An ima 1s)Animals Animals)
r SHUTTLE-TOUI{
OBSlRV/ITIOIJSL_
(Anima ls <25 m
L~from Ven)
0 1 (4)16 (64 )B (32)25
1-100 7 (41)3 (18)7 -(41)17
101-200 0 (0)0 (0)2 (100)2
,[201-400 1 (50)0 (0)1 (50)2
>400 0 (-)0 (-)0 (-)0
Total 9 19 18 46
n SHUTTLl-TOUR,
OHSERV fIT!OIlSt...-_._-----
(Animals >25 m,r from Veil)
'-..j 0 0 (-)0 (-)0 (-)0
1-100 4 (80)0 (0)1 (2U)5
101-200 27 (73)9 (24)1 (3)37
U 201-400 0 (-)0 (-)0 (-)0
>400 0 (0)5 (100)0 (0)5
Total 31 14 2 47
'-~-....
-'~SIIUTTLE-TOUR
OBSERVA TI or~s
(Total Animals)
[0 1 (4)16 (64 )8 (32)25
1-100 11 (50)3 (14 )8 (36)22
101-200 27 (69)9 (23)3 (8)39
201-400 1 (50)0 (0)1 (50)2
[~>400 0 (0)5 (100)0 (0)5
Total -40 33 20 93
[INTEHSIVE
OBSlRVA TI OI~S
AT UEN
r 0-100 47 (34 )70 (51)20 (15)137
101-200 67 (73)25 (27)1 (1)93
40U-GOO 48 (87)6 (11 )1 (2)55
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Total 162 101 22 iW5f-
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comments are made in the literature on flight distances for foxes.
Those reported generally describe animals that had apparently habitu-
ated to man.Several authors reported foxes ignoring the presence of
people 2-50 m away (Dixon 1938,A.Murie 1944,Rue 1969).
Relationshio of Reaction to Tvpe of Disturbance
Table 49 presents the actions of vehicles and visitors at active
fox dens near the road when faxes were visible.These data were col-
lected during the intensive observations at fox dens.At the Sable
Den,only about 50 m from the road,the public buses stopped more fre-
quently (68%)than private vehicles (54%),so that visitors could ob-
serve the foxes.Persons in private vehicles probably failed to sight
the animals as often as those on public buses,where more pairs of eyes
and a trained and experienced driver make sighting animals easier.Also,_
visibility of the den was better from the higher elevation of the bus
seats.Once the animals were sighted,however,and a vehicle stopped,
persons in private cars tended to get out of their vehicles more often
(36%)than persons in public buses (9%).For faxes at dens over 100 m
from the road,few persons spotted the animals and stopped their vehi-
cles.During the shuttle-tour observations,the bus passed 10%of the
foxes sighted without stopping;stopped,without people getting out for
76%of the sightinqs;and stopped,with people getting out for 14%of
the si ghti ngs .
The reactions of fox adults and pups within 200 m of the road to
various actions of vehicles and visitors,as I observed them during the
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Table 49.Actions of vehicles and visitors at red fox dens,1973-1974,l'!ount McKinley National
Park,{Ilaska.
ACTION OF VUIICLlS TYPE UF VElIICLE TOTAL
ANU VIS!TORS
NPS VEHICLE PUBLIC BUS PRIVATE VEIIICLE
Num5er (Percent lIumber (Percent iiumbel'(Percent Ilumber (Percent
Vehicles Vehicles)Vehicles Vehicles)Vehicles Vehicles)Vehicles Vehicles)
IJEIlS HlTflIrj
50 m OF ROAO
Pass >15 mph !l (80)14 (30)11 (42)33 (40)
Pass <15 mph 0 (0)1 (2)1 (4)2 (2)
Stop,people remain
in vehicle,quiet (10)29 (62)8 (31)38 (46)
Stop,people rewain
in vehicle,noisy (10)0 (0)(4)2 (2)
Stop,people out of
vehicle on road,
quiet 0 (0)0 (0)4 (15)4 (5)
Stop,people out of
vehicle 011 road,
noisy 0 (0)(2)0 (0)(1 )
Stop,people off
road,quiet 0 (0)2 (4)(4)3 (4)
Stop,people off
road,noisy 0 (0)0 (0)0 (0)0 (0)
Total 10 47 26 83
DEliS 50-200 m
FRO/I ROAU
Pass >15 mph 4 (100)23 (85)22 (92)49 (89)
Pass <15 mph 0 (0)0 (0)0 (0)0 (0)
Stop,people remain
(4)in vehicle,quiet 0 (0)4 (15)5 (9)
Stop,people remain
in vehicle,noisy 0 (0)0 (0)0 (0)0 (0)
Stop,people off vehi-
(4)cle,on road,quiet 0 (0)0 (0)(2)
Stop,people off vehi-
(0)(0)(0)(0)cle,on road,noisy 0 0 0 0
Total 4 27 24 55
176
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intensive den observations,appear in Table 50.A chi-squared analysis
of the reaction patterns exhibited tov/ard the various actions of the
vehicles and visitors reveals a significant difference (X 2=9.8,P<O.Ol)
betvleen the responses to a vehicle passing as compared to a vehicle
stopping,with the people remaining in the vehicle.The percentage of
mild and strong responses increased when a vehicle stopped.No signi-
ficant difference in responses was found between responses to a vehi-
cle stopping,with the people remaining in the vehicle,as compared
to a vehicle stopping,with people getting out.The sample size for
this comparison was small.
Relationship of Reaction to Aae and Sex
During 140 hours of intensive observations at one fox den within
50 m of the road,108 responses of pups and 28 responses of the adult
female to vehicles and visitors were observed.A chi-squared test
showed no significant difference in the reaction patterns (none:mild:
strong)of the pups compared to the adult female,although some behavior
patterns differed as described below.The adult female at this den
appeared thoroughly habituated to vehicles and visitors and exhibited
no,or only mild,responses to vehicles and persons on the road.The
adult male from this den was observed at the den on only four occasions,
all at night.He appeared much more wary than the female and pups.
The adult male was the only member of the family that frequently
watched the parked vehicle from which I made observations and circled
well around the vehicle when passing by it.On only one of the male's
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Table 50.Reactions of red foxes,adults and pups at dens within 200 m of the road,to vehicles
and visitors,1973-"1974,Mount HcKinley National Park,Alaska.
ACTION OF REACTION OF RED FOXES TOTAL
VEHICLE NU~lBER
ANU VISITORS NONE MILD STRONG AN Ij.jALS
Number (Percent Number (Percent Number (Percent
Anima 1s Animals)Animals Animals)Animals Animals)
Pass >15 mph 69 (60)43 (37)4 (3)116
Pass <15 mph 0 (0)3 (75)1 (25)4
Stop,people remain
in vehicle,quiet 34 (40)38 (45)12 (15)84
Stop,people remain
in vehicle,noisy 0 (0)2 (100)0 (0)2
Stop,people out of
vehicle on road,quiet 4 (40)5 (50)1 (10)10
Stop,people out of
vehicle on road,noisy 0 (0)1 (l00)0 (0)1
Stop,people walk
off road,quiet 6 (50)3 (25)3 (25)12
Stop,people walk
off road,noisy 0 (-)0 (-)0 (-)0
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visits-aid a vehicle drive up and stop near the den.The male respond-
ed by leaving the area immediately.
I observed another den over 800 m from,and out of view of,the
road.The den was observed from a distance of 75 m,which was as far
away as an observer could get and still see it.Again,the adult male
was the most disturbed by my presence.He usually barked for several
minutes,then left the denning area,when I was present.
At the other dens observed,not enough responses of the two adults
were observed to distinguish differences.However,one male regularly
rested during the day at a den 500 m from,and in view of,the road.
At this distance,the male occasionally raised his head to watch pass-
ing vehicles.
Allison (1971),A.Murie (1961),and Rue (1969)conmented that
male foxes often appear more wary than females.However,Rue (1969)
observed a den in McKinley National Park for several days and noted
that the male was much more tolerant of human presence than the female.
Use of Road as Travel Route
On many occasions foxes were observed traveling along the road.
They may be attracted to the road as an easy travel route.Usually the
distances traveled on the road were not known,but distances of up to
2 km were recorded.During 23 visits to a den within 50 m of the road
by the adult female,she came to the den a long the road on 10 vi sits
and left along the road on 9 visits.When the pups from this den began
venturing away from the den,they often left and returned along the
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road.Pups from three dens were known to occasionally use the road as
a play area.Adults were also observed hunting in ditches and on road-
sides as they traveled alonq the road.On one occasion an adult fox
captur~d a ground squirrel which had been temporarily distracted by a
passing vehicle.A.Murie (1944)and Rue (1969)described faxes hunt-
ing along the road in the park.
Behavior Patterns Related to Disturbance
Allison (1971)described many of the.behavior patterns faxes ex-
hibit when disturbed.Table 51 summarizes the disturbance behavior
patterns I observed during intensive observations at one den within
50 m of the road.At this den the animals showed no visible response
to 50%of the vehicular and visitor activities along the road,and only
watched during 37%of the disturbances.During 2%of the distubances,
pups ran up to a den hole and then stood or sat and observed the human
activities.During 8%of the disturbances,pups entered the den and,
thus,were no longer in view from the road.The adult male tended to
leave the den site during disturbances.The adult female often ignored
human activities.She barked,causing the pups to 'enter the den,on
only one occasion,when a bus stopped with people remaining on the bus.
At a den 800 m from the road,that I visited several times on foot,the
male often barked persistently at me,while the female did not bark
except when the male was present and barking.Both adults investigated
me by circling and approaching to within 3 m.In August the pups some-
times barked when disturbed.
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Table 51.Maximum behavioral reaction of red foxes at dens "Jithin
200 m of the road to vehicles and visitors,as observed
during intensive observations,1973-1974,Mount McKinley
National Park,Alaska.
RESPONSE TO VEHICLES NU~lBER (PERCENT
AND VISITORS ANIHALS ANIMALS)
Ho Visible Response 113 50
~Jatch 85 37
~Jal k Away from Road 0 0
Trot Away from Road 1 <1
Run Away from Road 2 1
Run to Den Hole 5 2
Enter Den 17 8
Leave Den Area 3 1
TOTAL 226
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Barking by the adult serves as a warning of danger to the pups.
Allison (1971)found that pups under 6 weeks of aqe immediately entered
the den when an adult barked.Older pups often responded to a bark by
running to the nearest den entrance and waiting for the danger to pass.
Allison also described a IIhacking cough ll I'Jhich was apparently a more
emphatic danger signal.,causing even older pups to react immediately.
Foxes encountered away from den sites during shuttle-tour trips
showed a greater tendency to run from the road than is indicated by the
intensive observations at dens.Thirty-nine percent of these foxes
ran.Probably many of the foxes encountered away from dens are indi-
viduals that have less contact with,and are less habituated to,road-
related human activities than those individuals denning near the road .
Feeding of Foxes bv Visitors u _
During my study I was aware of only one fox,an adult,that regu-
larly begged for food along the park road.This fox was often seen
along the road east of Savage River,where the road was open to unre-
strained use by private vehicles.The fox was observed traveling along
the road and trotting up to vehicles that slowed or stopped,often being
rewarded with food from the visitors.The adults and pups at the den
within 50 m of the road,where I made many of my intensive observations,
were never observed begging or being fed.This den is located along
the section of road where private vehicle use has been restricted since
1972.HOl'/ever,a long-time resident,Charlie Ott (pers.comm.1974),
remembers past years when pups using this same den had become beggars,
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running to the road when vehicles passed.Perhaps restricting private
vehicle use has reduced the amount of feeding of foxes by visitors.
Summary and General Discussion
Red foxes were observed on 63%of the shuttle-tour trips in 1973
and on 34%of the trips in 1974.Nine families in 1973 and four fami-
lies in 1974 were known to use dens within 400 m of the road.
No evidence was found that foxes,in general,avoid the vicinity
of the road during daily activities or during denning.Son~individ-
uals may have been attracted to the road.The road was frequently
used as a travel route by foxes.One fox was frequently fed by visi-
tors and purposefully approached vehicles.
Nany of the foxes denning along the road I'Jere habituated to human
activities.Adu1 t males seemed more wary than females.They often
left the denning area in response to human activities.Because of
small sample sizes,these response differences may have been related
to individual,rather than sexual,differences.In either case,if
differences cause one adult of a pair to make fewer visits to the den
with food,the survival of the pups,or energy demands on the other
adult could be adversely affected.
Stopping a vehicle increased the frequency and strength of re-
sponses significantly compared to those elicited when vehicles just
passed.
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Other r''1amma 1s
Wolf (Canis lupus)
The wolves of Mount McKinley National Park,as elsewhere,have
attracted the interest of researchers (A.Murie 1944,Haber 1968,
1972a)and visitors alike.The number of wolves in the park,east of
Wonder Lake,has been about 40-50 in recent years (Haber 1972a,1972b).
They are distributed throughout the park.Chapman (in prep.)is cur-
rently prepari ng a deta i1 ed thesi s on the effects of human di sturbance
on wolves in McKinley Park and elsewhere.Because this work will SOon
be available and because my own observations are limited,my comments
here are bri ef.
Wolves are observed much less frequently than the other large
mammals already discussed.During the 70 shuttle-tour trips,wolves
were observed on 5 occasions (7%of the trips).Each sighting was of
a single animal.Two of the animals ran across the road in front of
the approaching bus.The other three were each 300-400 m from the road
when sighted.All the wolves sighted on these trips exhibited strong
reactions to the buses,running away from the road and out of view.
However,wolves near the road did not always react strongly to vehicles,
as discussed below.During my plot surveys I observed wolves on three
occasions,once in the Sable Road Plot and twice in the Highway Off-
Road Plot,which was within 2 km of an active wolf den.
Besides the sightings mentioned above,during my general activities
in the park I sighted wolves near the road on 10 occasions.During four.
of these sightings the animals did not,at least immediately,react
strongly to the presence of one or more vehicles.All the animals
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involved were more than one year old.One wolf was observed for 7 min
as it traveled and hunted,covering about 1.5 km,at distances of
100-300 m from the road.lvly vehic 1e and a tour bus stopped to observe
the animal.The wolf glanced in the dire~tion of the road,but not
directly at the vehicles,several times.It successfully captured an
arctic ground squirrel and lay down to eat before continuing on.It
was finally lost from view behind a hill.On another occasion,two
wolves were observed resting 500 m from the road without showing
visible reactions to passing vehicles.On a third occasion,a wolf
was approaching the road at 20 m when I rounded a corner in my truck,
sighted it,and stopped.The wolf whirled,ran 3 m,then stopped and
began sniffing at the ground and investigating holes,as if hunting.
It proceeded in this manner,never again looking at the truck,until
it was lost from view 10 min later 300 m from the road.On the fourth
occasion,I rounded a bend in the road and observed a black wolf stand-
ing in the road 20 m ahead.I stopped my truck.The wolf continued
standing,looked at the truck for about 10 sec,then slowly approached
the truck,moving off the road about 2 m.At this time a shuttle bus
approached and stopped immediately behind my truck.A man with a
camera got out of the bus and quickly approached the wolf to within
10 m.The wolf watched the man and walk-trotted out of view into a
small ravine 30 m from the road.It reappeared 40 m from the road,
stopped,stared at the man,sniffed at the ground three times,and
then walk-trotted another 20 m away and disappeared into another ra-
vine.It was rumored among seasonal rangers that a wolf in the area
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of this sighting had been fed on one or more occasions by visitors,
but I did not talk to anyone who had actually seen this happen.It is
interesting that the animal approached my vehicle after I stopped.No
wolves in McKinley have been known to become real beggars.However,
experiences along the trans-Alaskan pipeline route have shown that
wolves can become habitual beggars where feeding by humans is not con-
trolled (Dan Roby,pers.comm.1976).
Wolves,like the red fox and other species,sometimes use the park
road as a travel route.On one occasion I observed a wolf travel 4.5
km along the park road in 40 min.A.Hurie (1944)observed tracks in-
dicating that wolves at times use the road in winter as a route for
easy access to Dall sheep on Polychrome Pass,and were even able to
kill sheep that used the road as a place to rest.Many authors have
provided evidence of the use of roads,human hiking trails,and snow-
machine or snowshoe trails as travel routes by wolves (Mech 1970,Peters
1974,Peterson and Allen 1974).Peters (1974),in an extensive study
of travel patterns of wolves in Minnesota,concluded that although
wolves used human roads and trails,especially in the winter,these
did not cause great alterations in their natural travel patterns.R.
O.Peterson and Allen (1974)reported a 90%decrease in wolf scats
found along hiking trails in Isle Royale National Park two weeks after
human visitation began in the spring.
During the period of my study,wolves were observed scavenging
one moose that had been killed by a vehicle.Wolves returned repeat-
edly to the carcass despite the fact that it was lying in the road
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ditch.I observed single wolves on the carcass on two occasions;each
time they ran into cover \'/hen vehicles approached.Large manJlIals are
only rarely killed by vehicles on the park road and the scavenging of
road-kill carcasses could not be an important food source for wolves.
Carbyn (1974a,1974b)commented on road-kill scavenging by wolves in
Jasper National Park,Alberta.
Wolf kills of large mamilals are not frequently observed from the
park road.During the period of my study persons in a tour bus ob-
served wolves chasing and killing a caribou 150 m from the nearest
point on the road,but 500 m from the point of observation.This car-
cass was completely utilized,first by wolves,especially at night,
and later by grizzly bears,over a period of several days.Richard
Chapman (pers.con~.1976)observed one wolf feeding at the carcass at
night as a vehicle with its lights on crossed a bridge 500 m away.The
wolf stopped,watched the vehicle momentarily,and then continued feed-
ing.On another night a different wolf,attempting to approach the
carcass,ran towards cover when a vehicle crossed the bridge.On an-
other occasion,wolves attacked a caribou on a river bar within 75 m
of the park road,but were apparently frightened off by a private ve-
hicle that arrived during the attack (Earl Sinns,pers.conID.1973).
The badly wounded caribou ren~ined at the same spot and was killed by
a brown bear later the same day.Road-related disturbances to wolves
at carcasses or during attacks appear to be infrequent.
The effects of the road on the denning of wolves are not well
documented.Since 1973 active wolf dens have been protected by special
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closed area status,which prohibits people from approaching the dens
closely.As with foxes,the wolves choose their whelping dens and bear
their pups before the road opens to vehicular traffic in the spring.
In 1973,wolves used a den only 100 m from the road for at least blo
weeks after the road VIas opened to traffic.They then moved the pups
to a den about SOD m from the road,and mostly hidden from view,where
they remained until the end of the denning period.In 1974,a wolf
family occupied a den 2 km from,and in view of,the road during June
and part of July.From this den a long stretch of road on Polychrome
Nountain is visible.Richard Chapman (pers.comm.1976)observed this
den for 335 hours and found that the wolves generally ignored vehicles
passing on the road,although he once observed two adults watching a
particularly noisy bus ascending the road up Polychrome Mountain.
These observations illustrate that wolves may successfully den in close
proximity to the road,at least if persons are kept from approaching
the den.Individual wolves undoubtedly differ in their tolerance to
the road (Chapman,in prep.).Carbyn (1974a,1974b)discussed the
abandonment of wolf dens in areas of heavy human use in Jasper National
Park.A.Murie (1944)described an extreme case of disturbance,during
which he entered the den and took a pup,after which the adults and
remaining pups continued to use the den.
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o
Lynx {Lynx canadensis}
Lynx were not frequently sighted in the park in 1973 or 1974.
During the 70 shuttle-tour trips lynx were sighted on two occasions,
one single and one pair.During my plot surveys I observed lynx once,
an adult female and three kittens,in the Igloo Road Plot.Thesingle
lynx sighted from a bus sat 3 m from the road for 5 min while visitors
observed it from the bus.It then walked para 11 e1 to the road and out
of view.The pair sighted from a bus was seen only briefly walking in
a ravine 100 m from the road before they were out of view.I sighted
the female and three kittens while I was walking 200 m off the road.
Although the female was aware of my presence,she allowed me.to sit
in full view and observe the family from less than 100 m away for
20 min.The female roamed over a small clearing,then lay dO\'Jn while
the kittens played and explored.Finally,the family wandered into
the shrubs and out of view.During this time several vehicles passed
on the road 200 m a\'/ay.The female occas ionally 91 anced to\'Jards them.
Wolverine {GuZo guZo}
Wolverines were not sighted during any of the shuttle-tour trips
or plot surveys.During 1973 and 1974,I observed one wolverine
traveling 350 m from the road.It immediately ran out of view as my
vehicle approached.
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Porcupine (Erethizon dorsatum)
Forty-six porcupines were sighted during the 70 shuttle-tour trips.
They \'lere observed on 2n of the trips.Fifty percent of the pOl~cupines
sighted were on the road,especially the vegetated edges of the road.
Table 52 shows the distances from the road for all porcupines observed.
All observations were of single animals along the first 55 km of the
road.Thirty-three percent of.the porcupines observed were in spruce
woods,28%in dwarf shrub tundra,24%in mixed open spruce and dwarf
shrubs,13%on gravel bars and 2%in tall willows.Buses often did
not stop to observe a porcupine,if one had already been sighted earli-
er during the same trip.The-buses passed,without stopping,54%of
the porcupines;they stopped,without people getting out,to observe
45%of the animals;and they stopped,with people getting out,for 4%
of the animals.Nearly all the porcupines that were on the ground
within 20 m of the road hurried away when the bus passed oor stopped.
The road appears to have an influence on the distribution of por-
cupines in the early summer.During both the shuttle-tour trips and
my road census trips in the research vehicle,porcupines were sighted
frequently before June 15 and rarely after this date (Table 53).This
trend was apparent in both 1973 and 1974 although these animals were
more numerous in 1973.Porcupines appear to be attracted to feed on
the young green vegetation on the roadsides in late May and early June.
As discussed earlier under IIBrown Bears",the first nevI qreen vegetation
of the year in a given area usually arpears on the roadsides.After
Observations of porcupines during shuttle-tour trips,
1973-1974,Nount r~lcKinley National Park,Alaska.
DISTANCE FRm'l ROAD (m)
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Table 52.
Number of
Porcupines
Observed
o
23
1-5
9
6-10
6
11-50
6
51-100
2
TOTAL
46
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Table 53.Numbers of porcupines and snowshoe hares observed along the road before and after
June 15,1973-1974,Mount McKinley National Park,Alaska.
AVERAGE NUMBER"OF
TYPE OF OBSERVATION NUf\1BER OF AVERAGE I'JU~1BER OF SNOWSHOE HARES
DATE TRIP TRIPS PORCUPINES PER TRIP PER TRIP
1973
Before June 15 Shuttle-Tour 9 3.22
After June 15 Shuttle-Tour 32 0.22
Before June 15 Research Vehicle Road Census 7 6.43 24.86
After June 15 Research Vehicle Road Census 10 0.30 2.20
1974
Before June 15 Shuttle-Tour 6 0.83 2.83
After June 15 Shuttle-Tour 23 0.22 0.65
Before June 15 Research Vehicle Road Census 5 1.40 8.20
After June 15 Research Vehicle Road Census 8 0.12 0
......
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new vegetation is widely available,porcupi~es are rarely seen feeding
on the roadside.
Snowshoe Hare (Lepus ameY'icanus)
Snowshoe hares were abundant during the early summer of 1973,but
declined in numbers from that time through 1974 (Table 53).In 1973,
hares were not recorded during the shuttle-tour trips because keeping
records on the large number of hares interfered with observing large
mammals.During the 1974 shuttle-tour trips and my research vehicle
road census trips both years,records were kept on the hares.The
average number observed per trip appears in Table 53.This table shows
that hares,like porcupines,were observed on the roadsides much more
frequently in the early summer,when the first green vegetation ap~
peared,than later in the summer.They were apparently attracted to
the roadside vegetation.Nearly all the hares were observed along the
first 56 km of the road,in spruce woods,deciduous woods,and dwarf
shrub habitats.Buses seldom stopped to allow visitors to observe snow-
shoe hares,and the hares almost always ran to cover when vehicles passed.
Hares are killed on the park road with some regularity by vehicles.
During 17 road censuses in 1973,eight road-kill hares were counted.
During 13 similar trips in 1974,when hares were scarce,no road-kills
were seen.
Besides being attracted to the greening vegetation on the roadsides
in early summer,hares were also attracted to two short sections of the
road throughout the summer.Along the first 2.4 km of the road,a wide
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section of disturbed ground at the side of the road has been stabilized
with a rich mixture of grasses and herbs.Snowshoe hares were frequent-
ly observed feeding in concentrations on this vegetation,especially at
night.During the research vehicle road censuses,15%of the hares
sighted were seen along this stretch of road.Hares were also frequent-
ly observed on the road along a 1.5 km stretch just west of Savage
River.Eighteen percent of the hares sighted during research vehicle
road censuses were observed along this short stretch of road,while 44%
of the hares sighted from the shuttle-tour buses in 1974 were observed
here.These aninals frequently appeared to be licking dirt from the
road along this stretch.It is possible that special minerals are
available here.Dale Guthrie (pers.conm.1976)has found gravel in
hare pellets in this area.Short-eared owls were also observed concen-
trating along this stretch of road and may have been attracted by the
hares.
Marmots (Marmota caligata)
An average of 1.3 marnnts per trip were observed on the shuttle-
tour trips.Marmots were sighted on 44%of these bus trips.All the
observations,except one,were made on Polychrome Mountain.Sixty-six
percent of the animals sighted were within 20 m of the road.The buses
stopped for the visitors to observe the marmots 85%of the time.
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Arctic Ground Squirrel (§?ermophiZus unduZatus)
Arctic ground squirrels are abundant in the park.They are occa-
sionally killed by vehicles on the road.Four road-killed squirrels
were counted during the 30 research vehicle road census trips.Several
species of wildlife,especially common ravens,magpies,mev;I gulls and
long-tailed jaegers,were seen feeding on road-killed squirrels.Ground
squirrels are frequently fed by visitors,despite regulations prohibit-
ing feeding.Beggar squirrels occurred at several points along the
road,including the hotel,campgrounds,visitor's center,and major
'scenic viewpoints.
Collared Pika (Ochotona coZZaris)
Pikas are found where suitable habitat occurs along the road.The
building of the road has created some new habitat.Pikas live in road-
side scree slopes in Igloo Canyon and on Polychrome Mountain.
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Short-eared Owl (Asia llammeus)
In 1973,when microtines were relatively abundant,an average of
2.4 short-eared owls per trip were sighted on the shuttle-tour trips.
They were seen on 71%of the trips.In 1974,with microtines reduced
in numbers,only 0.14 owls per trip were sighted,with the birds being
observed on only 14%of the trips.The buses stopped so that the visi-
tors could observe the owls during only 31%of the sightings.Short-
eared owls appeared to be attracted to the road as a hunting area ..
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Microtines,arctic ground squirrels,and snowshoe hares crossing or
wandering on the road may be particularly easy prey.In 1973,during
periods of twilight and darkness,I flushed as many as 30 short-eared
owls from the road or roadside,while driving the first 50 km of the
road.Owl s vlere particul arly concentrated on the road,with as many
as eight observed at one time,along a 1.5 km stretch just west of
Savage River.Snowshoe hares,perhaps seeking special minerals,were
often seen on this section of road and their presence may have attract-
ed the ov/l s.
Long-tailed Jaegers (stercorarius Zongicaudus)
An average of 0.9 long-tailed jaegers were sighted during each of
the shuttle-tour trips.Jaegers were observed on 44%of the trips.
These birds attracted the attention of visitors more than most other
bird species.The buses stopped during 74%of the sightings.I was
aware of three jaeger nests within 100 m of the road each summer.One
of these nests vias known to have hatched,although the fate of the
others is not known.Just after fledging,a young jaeger from the
successful nest was observed on several occasions feeding on road-
killed ground squirrels near the nest site.One parent was also seen
feeding on road-killed squirrels.These birds allowed vehicles to
approach vlithin 5 m vlhile they \vere on the road feeding.
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Willow Ptal~igan (LapoDus lagopus)
Records were not kept on ptarmigan during the shuttle-tour trips.
Notes were made on ptarmigan observed during the research vehicle cen-
sus trips.The numbers of ptarmigan observed during these trips appear
in Table 54.Ptarmigan were more abundant in 1973 than in 1974.In
early spring,males often used the raised roadbed as a vantage point
from which to survey their territories (Table 54).They also used
roadside willows as vantage points.Many willows grow along the road,
because of drainage patterns from the road,while being absent from
nearby areas.Later in the summer I observed families feeding on the
roadside,especially where Eriophorum was localized in wet roadside
ditches.Ptarmigan were also observed taking gravel from the road.
Thus,these bi rds are attracted to the road for a val~i ety of reasons.
Males on the road in the spring were sometimes bold in their reac-
tions towards vehicles.During my road surveys,I drove slowly past
ptarmigan.They responded in the following manners:12%flew away from
the road,22%walked or ran a short distance off the road,and 65%
remained on the road,usually crouching low and watching my vehicle
pass.A few males ran aggressively towards my vehicle or ran along
in front of it,as if leading the vehicle out of their territories.
t~ew Gull (Larus canus)
During my road census trips in the research vehicle,an average
of 9.5 mew gulls per trip were observed on the 12 spring trips.None
were observed on the fall trips.Mew gulls,like ground squirrels,are
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Table 54.Numbers of willow ptarmigan observed during road census
trips in research vehicle,1973-1974,Nount rlcKinley
National Park,Alaska.
DATE NUt·1BER AVERAGE NUMBER OF PTARMIGAN
OF PER TRIP
TRIPS
ON ROAD TOTAL OBSERVED
1973
~1ay 21 -Hay 24 6 12.00 32.67
June 21 1 3.00 4.00
August 30 -September 30 10 5.10 ·11.80
1974
Hay 25 -fvlay 29 2 2.00 14.00
Nay 30 -June 1 3 0 1.00
August 28 ~September 8 8 0 a
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frequently fed by visitors and some have become beggars.During the
shuttle-tour trips,up to 11 gulls were counted at the Eiclson Visitor
Center,feeding on visitor handouts.Mew gulls were frequently ob-
served flying along the road,following buses and other vehicles,and
twice landed on my vehicle to peer through the window.They also
frequent campgrounds and the garbage dump (Downing 1975).
200
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FINAL DISCUSSION
General Biological Concepts and the Responses of Wildlife to Human
Acti vi ti es
Many general concepts in the fields of animal behavior and ecology
are directly useful in understanding the responses of wild animals to
human activities.A key factor influencing the behaviQr of higher
vertebrates,especially mammals,is their ability to learn.Mammals
function best physiologically in a familiar,predictable environment
(Geist 1971a,1975).Maximum energy efficiency in the utilization of
their environment only occurs when sources of fear and uncertainty are
minimized (Baskin 1974,Cowan 1974,Geist 1975).Mammals often re-
spond to strange new situations or stimuli with a combination of alarm
and curi os i ty.Later,through 1earni ng,they modi fy thei r responses
appropriately to reduce uncertainties and danger.Depending on the
outcome of their first encounters with a new stimulus,they may learn
to avoid,approach or ignore it.Thus,the response of mammals to human
activities depends to a large degree on the actions of people during the
animals'first encounters with them.If alarm and fear responses are
not reinforced,habituation occurs.If the animals are rewarded during
encounters with humans,they may become attracted to people.Responses
learned during early experiences with a given stimulus may be general-
ized to other stimuli perceived as similar by the animal (Cowan 1974).
In national parks,the habituation of animals to common human
activities and artifacts,such as vehicles and roads,should be
201
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encouraged.Habituation allows wild animals to efficiently utilize
habitat near centers of humar.activity (Geist 1971a).It also pro-
vides visitors with an excellent opportunity to observe wild animals
in near-natural environments.Alternatively,if park animals are re-
peatedly frightened and learn to fear humans,they may avoid areas of
human activity or expend critical amounts of energy in physiological
stress and frequent fear responses.Both situations could adversely
affect the health and population dynamics of the species involved •
In order to take advantage of and encourage the capacity of mam-
mals to habituate to human activities,an understanding of the stimulus-
response mechanisms of the species in question is necessary ..Animals
continuously filter the stimuli coming to them from their environment
and selectively respond to key stimuli.The perception of a situation
and its accompanying stimuli by one species may be quite different than
the perception of the same situation by another species,including man
(von UexkUll 1957).The perceptive abilities and relative importance
of the various senses in the large mammal species found in r1cKinley
Park have never been studied in detail.However,previous discussions
in this report indicate that smell is of major importance as a danger
signal to caribou and brown bears,while hearing is important to moose
and sight is important to sheep.
The ability of an animal to habituate varies with the type of
stimulus to which it is subjected.Berqerud (1974)suggests that the
responses to certain stimuli have an ontogenetic origin and are not
easily modified by learning.These stimuli are usually key factors
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used by the species to identify dangers that have been present during
its evo1 uti onary hi story.Important evo 1uti onary factors i nf1 uenci ng
disturbance responses include predator-prey relationships and natural
environmental hazards.Stimuli of importance to an animal may seem
subtle to humans.Pruitt (1965)discovered that caribou responded
strongly to the outline of a man wearing a wolf-ruff hood around his
head,but did not respond to the same man when the hood was down.He
postulated that the tall,narrm<J outline,with a broadly rounded top,
of a man with a hood was very similar to the outline of a hunting wolf
approaching directly,with its head lowered and the broadly rounded
ruff of its shoulders providing the top outline.Non-hunting "'lo1ves
generally have their heads up and present a different outline.Klein
(1971)noted that reindeer refuse to cross bridges and suggested that
the hollow sound of a bridge produces the same stimuli to the reindeer
as thin ice.A common stimulus situation that may produce ontogenetic-
ally-based fear responses in a variety of species is a direct,close
approach,as this is often associated with predators.Thus,direct
approaches by park visitors reinforces the alarm response of a naive
animal and inhibits habituation to visitor activities.
Other types of stimuli,besides those releasing ontogenetically-
based fear responses,may generate alarm.These include stimuli that
are not a familiar or predictable part of the environment or stimuli
that generate sharp contrasts in the environment,such as loud noises
or sudden movements (Geist 1971a).To e~courage habituation these types
of stimu1 i shoul d be avoi ded duri ng encounters bet\'/een humans and wil d
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animals.
Besides predator-prey relationships,many factors in the life
strate~y of a species are important in understandinq the responses
of animals to human activities alonq a road.Home range size affects
the frequency of encounters between individuals and road-related acti-
vities,and the frequency of encounters affects the habituation process.
Habituation occurs only if a harmless situation becomes familiar and
predictable (Geist 1971a).Species with small home ranges which in-
cl ude the road woul d frequently encounter human acti viti es,a 11 o\A/i ng
habituation or the buildup of physiological stress,depending upon the
nature of the encounters.Individuals with large home ranges may
rarely encounter the road,resulting in less opportunity for habitu-
ation.The discussions in this report provide evidence that moose,
Dall sheep and brown bears,especially females with young,all have
restricted seasonal home ranges of only a few square kilometers that
are often used in successive years.Caribou tend to be more nomadic
than the other species.Male brown bears often utilize large home
ranges.Naive caribou and naive male brown bears are probably more
frequently sighted from the road than naive moose,sheep,or brown bear
families,and may be expected to be less habituated to human activities.
The large mammals in the park generally have relatively long life spans
of 10-20 years (r1undy and Flook 1973,A.t~urie 1941,R.L.Peterson
1955).For the species with restricted home ranges,the same indivi-
duals are often observed from the road throughout a summer and in suc-
cessive years.These individuals may also have been raised near the
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road.These individuals could easily become habituated to human acti-
vities if humans did not reinforce the animals'initial fear responses.
During certain critical periods in the annual cycle,the potential
damaging effects of disturbances are magnified.Disturbances during
periods of severe cold,deep crusted snow or food shortages in winter
can cause abortions,injuries,additional stress and death (Geist 1971a,
Zhigunov 1961).The park road is not open to vehicular traffic 9uring
the winter,so disturbances from human activities are greatly reduced
at this time.However,winter use by skiiers and mushers is increasing
rapidly.Calving time is also a critical period,since disturbances
can cause desertion,trampling and injury to the young (Zhigunov 1961).
Most of the caribou in the park calve away from the road,but very
young moose calves are frequently sighted from the road.Disturbances
during migration may delay or divert migrating animals,possibly affect-
ing their range use.In the park,sheep sometimes failed in attempts
to move from winter to summer ranges because they were disturbed while
attempting to cross the road.The large migrating bands of caribou
always remain south of the road,not attempting to cross.
Interacting Variables Affectinq Responses
A large number of internal and external variables interact in de-
termining the response of an animal to human activities.These factors
include distance and frequency of disturbance,type of human activity,
group size,sex,age,behavior before disturbance,motivation,social
facilitation,past experiences,individual temperament,health,time,
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206
season,weather and habitat.These factors have been discussed,by
species,"previously in this report.A few comparative or general com-
ments appear here.
Reaction Oistances
The response patterns (none:mild:strong)at ~iven distances were
remarkably similar for the four most fl~eouently observed larqe Jl1RP.1-
mals,i.e.caribou,moose,nall sheep and brown bears (Figs.6,9,
11,13).For caribou,moose and bears the distance at which 50%of
the animals exhibited strong reactions occurred beu~een 25 and 50 m
from the road.For Da 11 sheep thi s poi nt occurred betltJeen 10 and 25
m from the road.All four species passed the threshold of 50~&II no
visible reaction ll between 50 and 100 m from the road.All the species
exhibited 1ess than 10%strong responses beyond 200 m.There ItJere no
strong responses,except for three bear families,at distances beyond
400 m.Strong responses,by definition,included all flight behavior
except II VJa lking less than 10 m.1I Flushing distances are highly vari-
able between individuals and situations and mav be over 1 km under
certain conditions (de Vos 19GO,Jakimchuk et aZ.1974).HOI"!ever,
the average flushing distances in response to man reported in the lit-
erature all fall within a small range.For naive caribou,reported
averaqe flushing distances range from 90-160 m (Beroerud 1974,Kelsall
1957),for naive moose 80-14n m (Oenniston 1956,~1c~1illan 1954,Strinrjham
1974),for habituated moose 5-50 m (nenniston 1956,tld1illan 1954,
Stringham 1974),for habituated sheep 15-60 m (Anderson 1971,nixon
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1938,Jones et aZ.19(13),and for naive bears about 100 m (F.C.
Craigheaci and J.J.Crai~head 1%6).Thus,"t'lpical flushinq dis-
tances ll for these species seem to be 90-160 11 fr)l~naive animals and
5-60 m for habituated ani~n1s.
Some differences between species become apparent \'then one looks
at the distance in the shuttle-tour data at which 90%of the animals
exhibited no visible response to the human activities.This distance
was between 100-200 m for Dall sheep,200-300 m for broltJn bears,
300-400 m for moose and 400-500 m for caribou.Caribou showed the
highest percentage (41%of the animals)and sheep the lowest percent-
age (34%of the animals)of strong resnonses within 100 n of the road.
Caribou may be the most reactive species to road-related hUMan activi-
ties because they range over a 1araer area than the other species and
a hiqher percentage of naive individuals may be included in the encount-
ers.They are also highly social;one naive animal in a qroup can stim-
ulate responses by others.The results for Oall sheep sho\'ted them to
be particularly unresponsive to human activities.Few sheep occurred
within 200 m of the road because very little habitat for sheep ''las found
within this distance of th~road.Most observations of sheep close to
the road involved a small,well-habituated band on Polychrome Pass.
Brown bears were particularly unresponsive at distances beyond 200 m,
although their responses were more variable than other species.Adult
brown bears have f~1 enemies except other bears and man.Even man
has probably only been a serious predator since the development of
dependable firearms.Brown bears seem to exhibit fewer innate fear
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responses and less wariness than ungulates.The latter have evolved
under the.influence of effective predators.Many bears,especially
sows with families,remained in small areas near the road for long
periods and in successive years,thus having the opportunity to habi-
tuate.The great variability in responses by bears may result from
the fact that fewer responses are innate and responses are highly mod-
ified by individual experience.Bears encountered by the road a1so
include both highly habituated families and naive single animals which
range long distances and only occasionally encounter the road.
Type of Disturbance
The actions of the shuttle-tour buses varied slightly depending
on the species.The actions of the buses were as follows:1)they
passed,without stopping,2-12%of the individuals of each species
sighted,2)they stopped,with the people remaining in the bus,for
62-75%of the animals,3)they stopped,with the people getting out
but remaining on the road,for 13-32%of the animals,and 4)they
stopped,with people getting out and walking off the road,for 0-3%of
the animals.Tour buses tended to stop more often than shuttles,but
people tended to get out of shuttles more often than tour buses.
People on the buses seemed very efficient at sighting animals,both
because of the experienced drivers and because there were many pairs
of eyes searching in all directions.Observations of private vehicles,
although inconclusive,indicated that people got out of them slightly
more often and walked off the road to\!Jard animals more often than people
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in buses.People in private cars failed more often than the people in
buses to sight faxes at dens.The frequency of spotting animals from
private cars is not known for the other species.
For large mammals within 200 m of the road,only faxes showed a
significant increase in responses when vehicles stopped,with people
remaining in the vehicle,compared to passing without stopping.Since
buses infrequently passed animals that were Observed,useful sample
sizes often were not available.People getting out of the buses in-
creased strong reactions for caribou,moose and sheep but not for bears
or faxes (Figs.8,10, 12, 14,Table 50).It appears that the ungulates,
all of which have evolved under the influence of effective predators,
are more sensitive to the appearance of people than the carnivores.
The faxes involved were at dens near the road and highly habituated.
People from buses infrequently moved off the road but this action gen-
erally resulted in strong responses by the animals.As discussed earlier,
animals do not habituate quickly to the threatening stimuli of a direct
approach.Loud noises produced by shouting,whistling,honking,banging
on the side of the buses,screeching brakes and other means dramatically
increased strong responses from caribou,moose,sheep,bears and faxes
(Figs.8,10,12, 14,Table 50)."The increase was particularly great
for sheep and moose.Loud noises were effective in eliciting responses
probably because they provided sharp contrasts to the usual sound stimuli.
They were also unexpected,since they occurred durin9 only 10%of the en-
counters.Moose,which often inhabit dense vegetation,are particularly
dependent on hearing to warn them of danger (R.L.Peterson 1955).Sheep,
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.living in mountainous habitat t are vulnerable to rock slides and ava-
lanches and loud noises are of particular significance as a warning
signal to them (Dixon 1938 t Geist 1971c).
Sex and Aqe,
My study and the literature reviewed in the previous species dis-
cussions show that t in unhunted populations of caribou,moose and sheep,
females with young are generally the most sensitive to environmental
disturbances and old males are least sensitive.Reproductive success
for females depends on the protection of the young from predators.
Fleeing from sources of potential danger is a protective strategy used
by many ungulates,and females with young are often characterized by
long flushing distances.However,young moose calves need several days
to develop fully their locomotor abilities (Altmann 1958b).Female
moose with very young calves may have very short flight distances and
may attack a potential source of danger rather than flee (Altmann 1958b).
The males of these ungulate species depend on their size and condition
during the rut to become part of the reproducing population.By minimiz-
ing their responsiveness to environmental stimuli they would conserve
energy.If an attack occurs't a healthy male in his prime may more
easily escape predators than smaller,younger animals.
Effects of Disturbance
Geist (1971a,1975)reviewed the possible effects of various
levels of disturbance on large mammals.Minimally,escape behavior
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and anxi ety requi re the uti 1i zati on of energy.Thi s ener9Y mus t
either be replaced by increased food intake and thus increased pres-
sure on the food source,drawn from fat reserves,or redirected away
from other energy uses,such as grov/th,development,fat depos iti on,
and reproducti on.Energy costs depend on the i ntens i ty,dura ti on and
frequency of disturbances.Chronic excitation or stress can cause
physiological disorders (Zhigunov 1961).The mere act of fleeing,
especially panicky flight,may result in abortions,desertion or tramp-
ling of young (Zhigunov 1961).Repeated disturbances may cause abandon-
ment of habitat (Geist 1971c).All these responses may,in turn,ad-
versely affect reproduction and mortality.Unfortunately,field studies
of the actual energetic and physiologic costs of various types and levels
of disturbance have not been done.
My study did not reveal any large-scale adverse effects of human
activities on the wildlife near the park road.Many of the animals were
habituated to some degree to human activities,especially those confined
to the area of the road.Animals usually fled less than 200 m in response
to road-r2l~ted human activitiei.Longer flights occurred in a few
sensitive individuals,or when animals were persistently followed off
the road by visitors or chased along the road by a vehicle.The inten-
sive observations of caribou did show that during the day animals within
200 m of the road tended to mill around more and feed less than caribou
further from the road.These animals encountered human activity along
the road several times each hour and did not appear to completely return
to normal activities between encounters.Thus,the balance of energy
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intake versus energy expended was not as favorable for animals feeding
within 200 m of the road as for those feeding further away.This type
of relationship may have also been true for other species,but not
enough intensive observations were made on them to document it.
There is no evidence that the vicinity of the road was avoided by
a large number of individuals of any of the major species.A few sen-
sitive individuals probably did avoid the road.Male bears seemed to
be under-represented near the road.Fewer male bears may have been hab-
ituated to the road because their large home ranges resulted in each in-
dividual only infrequently encountering the road.
Ground squirrels and snowshoe hares,when abundant,were frequently
hit and killed by vehicles.Road-kills of large mammals were ve~y in-
frequent.
Attraction of Wildlife to the Road
A number of species were attracted to the road to feed on the road-
side vegetation.In the spring,the vegetation along the roadside began
its new growth before the vegetation in nearby areas,and porcupines,
snowshoe hares and,to a lesser extent,brown bears were attracted to
feed on this new growth.Throughout summer,the hares continued to
feed on grasses and herbs seeded to stabilize the roadside near the
entrance of the park.Hares were also attracted to one other short
section of the road,possibly by special minerals.Road drainage
patterns favored roadside sedges and willows in some areas.Moose
occasionally fed on the willows.Willow ptarmigan sometimes fed on
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the seeds of roadside sedges.They also obtained gravel from the road
and took dust baths on its surface.Male ptarmigan sat on the raised
road in the spring to survey their territories.
t1ew gulls,long-tailed jaegers,ravens and magpies \'/ere observed
feeding on squirrels and hares that had been killed by vehicles.Short-
eared owls were apparently attracted by the hares which concentrated
along parts of the road.Foxes frequently used the road as a travel
route and sometimes hunted along its edge.Ground squirrels and mew
gulls concentrated near centers of tourist activity,where they received
handouts of food from visitors.Some foxes were also observed being fed
by visitors.
Manaqement Recommendations
Many of the animals in t10unt tlcKinlev National Park are habituated
to various degrees to human activities along the park road.Habituation
should be encouraged,as it allows wild animals to utilize the roadside
habitat without expending large amounts of energy in fear responses and
physiological stress caused by human activities.Habituation of animals
also allows visitors to view the animals.At the same time,visitors
must be educated to respect wildlife and not to treat habituated wild
animals as domestic pets that can be approached closely,touched,
teased or fed.
To encourage habituation,stimuli that generate fear in wild ani-
mals should be avoided or minimized durinq human-wildlife encounters.
Such stimuli include direct approaches,"sneak"approaches,loud noises,
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quick movements,getting out of vehicles under certain circumstances and
other situations discussed previously.Animals habituate best to situ-
ations that are familiar and predictable.The more uniform the actions
of vehicles and visitors,the less disturbing they will be to the wild-
life.For instance,controlling the speed of vehicles helps minimize
disturbance.If most vehicles pass slowly and the animals are accus-
tomed to this action,then a vehicle speeding past usually stimulates
a strong response.Some uniformity of visitor activities has already
been introduced by the use of a public transportation system.This
system makes many of the'vehicles on the park road uniform in.appear-
ance and the activities of visitors on the public buses are more pre-
dictable than those of visitors in private cars.The presence of other
visitors and the bus driver inhibits 'blatant harassment or quick approaches
towards animals.The bus system also provides a plus for the park visitor
since the many pairs of searching eyes in a bus are usually able to spot
as much or more wil dl i fe than a we 11-tra i ned observer.
The drivers of the public buses are the park personnel with the
most contact with both visitors and wildlife along the road.They could
effectively moderate many visitor-wildlife encounters.Bus drivers should
be made to feel like an important part of the park system.During training,
bus drivers should be given a sound knowledge of and respect for the park's
wild animals.They should be made aware of the visitor activities that
are most disturbing to wild animals and which animals are most sensitive.
This knowledge would enable them to moderate visitor activities.Com-
munication of this information to passengers is equally important.This
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215
same information should be available to park visitors driving private
vehicles.
Animals in certain situations are particularly sensitive to dis-
turbance and the intensity of human interactions with animals in these
situations should be minimized,perhaps by having people observe quietly
from their vehicles.'Such situations include encounters with females
with newborn or small young,Dall sheep attempting to cross the road
while moving from one range to another,bears and wolves on carcasses,
wolves at dens and foxes at dens,if one or both of the adult foxes
appear sensitive to human activities.
Feeding of wildlife is already prohibited in national parks.
However,beggar animals still occur with some frequency.An infor-
mative explanation of the reasons for the "no feeding"regulation
should be made available to visitors and this regulation should be
strictly inforced.
The frequency of vehicles on the road during the day seems already
high enough to significantly affect the activity patterns of caribou,
and perhaps other species,within about 200 m of the road.Some of
these animals remain nervous for most or all of the time between en-
counters with vehicles and their feeding time is reduced.The fre-
quency of disturbances per visitor is reduced by using public buses
rather than private vehicles because more people can ride in each
vehicle.The frequency could be further reduced by encouraging vehicles
to travel together rather than spreading out a1on9 the road.
It would be useful to park managers to systematically monitor
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wildlife abundance and behavior along the park road.This could be
done using an observer on public buses,as in this study.Data from
such monitoring would provide early indications of changes in wildlife
abundance,distribution or behavior near the road.Such changes may
be correlated with changes in numbers of visitors,changes in visitor
use patterns,changes in management policies,or changes within the
wildlife populations or habitats unrelated to human activities inside
the park.
Much more research is needed on the physiological effects of various
types of human disturbances on wild animals.Disturbances could have
many serious,but not immediately visible,effects that cannot be de-
tected by a study such as the one reported here.Park managers could
benefit by encouraging and supporting species-specific research into
these problems,even if the work is not conducted within the park.
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APPENDIX A.SUPPLEMENTARY DATA TABLES
217
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Table Al.Dates of plot observations,Mount McKinley National Park,Alaska.
PLOT DATE OF OBSERVATION PERIOD
1 2 3 4 5
Igloo Road 1973 June 25 July 20 August 14 August 20 September 1
Igloo Road 1974 June 20 July 8 July 30 August 12 August 22
Igloo Off-Road 1973 July 1 July 9 August 10 August 18 August 26
Igloo Off-Road 1974 June 21 July 25 August 2 August 14 August 25
Highway Road 1973 July 4 July 23 July 30 August 15 August 23
Highway Off-Road 1973 July 21 July 28 August 9 August 17 August 24
Stony Road 1973 June 19 July 5 July 22 August 19 September 7
Stony Off-Road 1973 June 22 July 6 July 25 August 19 September 4
Sable Road 1973 June 30 July 22 July 27 August 15 August 25
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Table 112..Actual numbers and minimum densities of caribou observed during each plot observation period,first-search,
Mount McKinley National Park,Alaska.
PLOT OBScRVATIO/l PcRIOIJ (F1I{ST-SEAA~H)
2 3 4 5
Aninlills Groups Anir.1iI1s Groups Aninlills Groups An i filii 1s Groups Animals Groups
Dens ity Density Uensity Uensity Densi ty Uensity Uensi ty Density Uensi ty Uens.i ty
No.{Per kr,h flo.(Per km2 )flo.{Per knh flo.(Per knh flo.(Per knh No.{Per knh No.(Per kn/)No.(Per ki)No.(Per lui)No.(Per ki)
Igloo Road 1973 2 0.131 2 0.131 0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0
Igloo Road 1974 0.065 0.065 0.065 0.065 2 0.131 0.065 0.065 0.065 0.065 0.065
Igloo Off-Road 1973 2 0.104 0.052 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Igloo Off-Road 1974 0.052 0.052 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Highway Road 1973 4 0.223 2 0.112 0.056 0.056 2 0.112 2 0.112 0 0 0 a 0.056 0.056
Hig~way Off-Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 a
Stony Road 1973 0.081 0.081 0 0 0 0 8 0.642 2 0.161 0 0 0 a 0 0 0 0
Stony Off-Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 8 0.816 6 0.612 0 0 0 a
Sa~le Road 1973 10 0.482 2 0.097 13 0.628 6 0.290 6 0.290 3 0.145 2 0.097 2 0.097 0.048 0.048
,h 'li.j d',\;/r'"C:~(--[7"1 L:l c-J en l ~.r--~(.---'r-J I~r-J C__l r----]i-Y C~J l..",l ..J
Table AJ.Actual nu~ers and mininlUffi densities of caribou observed during each plot observation period.total day.
Mount ~kKinley National Park.Alaska.
PLOT OBSERVATION PERIOu (TOTAL OAY)
2 3 4 5
Anima 1s Groups Animals Groups Animals Groups AninJd1s Groups Animals Grcu?s
Density Density Density Oens i ty Dens i ty Uensity Uensity Uens i ty i.>ensity Dens i ti
/lo.(Per kroh /lo.(Per kJn2)No.(Per km2 )No.(Per ki)flo.(Per kr.h /lo.(Per km2)No.(Per kJn2)No.(Per kJn2)flo.(Fer ki)lio.(Per lu:iZ)
Igloo Road 1973 2 0.131 2 0.131 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 U
Igloo Road 1974 0.065 0.065 0.065 0.065 2 0.131 0.065 2 0.131 2 0.131 0.065 0.065
Ig100 Off-Road 1973 2 0.104 0.052 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 il
Igloo Off-Road 1974 0.052 0.052 0 0
0 0 0 0 0
0 0 0 0
0 0 0 0 0
Highway Road 1973 4 0.223 2 0.112 0.056 0.056 5 0.279 3 0.168 0 0 0 0 0.056 O.u56
Highway Off-Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Stony Road 1973 0.081 0.01l1 0.081 0 0 9 0.126 3 0.242 0 0 0 0
0 0 0 a
Stony Off-Road 1973 0 0 0 0 0 0 0 0 0 0 0 0
11 1.122 7 0.714 0 0 0 0
Sable Road 1973 10 0.483 2 0.097 15 0.725 8 0.386 11 0.531 4 0.193 2 0.097 2 0.097 2 0.097 2 0.097
N
No
r-J ~,l ,,'r--"'1'IL.:)
I;
lj
,
1-lJ
T~ble A4.Actual numbers and minimunl densities of ~se observed during e~th plot observatioq period,first-searth,
Mount McKinley National Park,Alask~.
PLOT (;~SEl\VATl(Hl PERIOD (FIRST -SI:.ARCIl)
2 3 4 5
Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups
Density Density DensHy lJensity lJensity lJensity lJensity lJensity liens i ty Dens ity
No.(Per km2 )No.(Per kr.h No.(Per ki)No.(Per ki)/lo.(Per kr,h /lo.(Per km2 )/lo.(Per ki)/lo.(Per kID2 )/lo.i:(Per kIDZ)(Per km )110.
Igloo Road 1973 2 0.131 0.065 0 0 0 0 3 0.196 Z 0.131 0.065 0.065 0.065 0.065
Igloo Road 1974 4 0.261 3 0.196 2 0.131 2 0.131 2 0.131 2 0.131 3 0.196 Z 0.131 3 0.196 2 0.131
19100 Off-Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0.052 0.052 2 0.104 2 0.104
19100 Off-Road 1974 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.052 0.052
Hi gr.way Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.056 0.056
Hi9~ay Off-Road 1973 0.078 0.078 0 0 0 0 0 0 0 0 0 0 0 0 0.078 (j.078
Stony Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Stony Off-Road 1973 0.102 0.102 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Saole Road 1973 0.048 0.048 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
N
N
....J
Table AS.Actual numbers and minimum densities of moose observed during each plot observation period,total day,
Mount ~IcKinley Uational Park,Alaska.
PlOT OIlSEkVATIOII PlRIOU (TOTAL UAY)
2 3 4 5
Animals Groups Anima Is Groups Anir:l3ls Groulis Anirll3ls Groups Animls Groups
Uensity Oensity Uens i ty Uensity Uensity Uensi ty Density Dens i ty Uens i ty Density
No.(Per ki)/.0.{Per knh No.{Per knh /10.(Per km2)No.(Per km2)/lo.(Per K1lI2 )Ho.{Per knh No.2 (Per km2 )No.(Per'~2)(Per km )iiO.
Igloo Road 1973 2 0.131 0.065 2 0.131 0.065 3 0.196 2 0.131 0.065 0.065 0.065 0.065
Igloo Road 1974 4 0.261 3 0.196 2 0.131 2 0.131 2 0.131 2 0.131 3 0.196 2 0.131 3 0.196 2 Ii.J:;I
Igloo Off-Road 1973 0 0 0 0 0 0 0 0 0 0 0 0
0.052 0.052 2 0.104 <:v.l04
Igloo Off-Road 1974 0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0.052 v.052
Highway Road 1973 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0.0:'6 U.056
Highway Off-Road 1973 0.078 0.078 0 0
0 0 0 0 0 0 . 0 0 0 0 0.078 0.078
Stony Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 u U
Stony Off-Road 1973 0.102 0.102 0 0 0 0 0 0 0
0 0 0 0 0 0 0 U u
Sable Road 1973 0.048 0.048 0 0 0 0
0 0
'0 0 0 0 0 0 0 0 0 U
N
N
N
_________________---,J
r '-[:-:J ,-----,
,J
Table M.Actual numbers and minimum densities of Call sheep observed during each plot observation period,first-search,
Haunt McKinley national Park,Alaska.
PLOT OBSERVA TlOIl PERWU (F IRST -SEARl-H)
2 3 4 5
AniNls Groups Animals Groups Animals Groups Animal s Groups Animals Groups
Uensity Density llensity llensity LJensity LJensity Uensity llensi ty Density Density
no.(Per ki)No.(Per krn2)flo.(Per ki)No.(Perki)flo.(Per km2)No.(Per krn2)110.(Per km')No.(Per ki)lIo.(Per km2)Ho.(Per Km2)
Igloo Road 1973 b2 4.052 9 0.5&1 0.458 2 0.131 26 1.699 2 0.131 32 2.092 3 0.196 14 0.915 0.065
19100 Road 1974 9 0.5!l8 5 0.327 27 1.765 4 0.261 15 0.980 2 0.131 17 1.111 2 0.131 1~0.9aO ,0.131
19100 Off-Road 1973 40 2.083 3 0.156 81 4.219 6 0.312 45 2.344 8 0.417 15 0.781 3 0.156 102 5.312 4 0.208
19100 Off-Road 1974 30 1.562 7 0.365 3 0.156 0.520 62 3.229 6 0.312 48 2.500 6 0.312 83 4.323 3 0.156
Highway Road 1973 8 0.447 2 0.112 9 0.503 0.056 3 0.16!l 0.056 0 0 0 0
0 0 0 0
Hishway Off-Road 1973 0 0 0 0 14 1.085 2 0.155 0 0 0 0 0 0 0 0 4 0.310 0.078
Stony Road 1973 0 0 0 0 a 0 0 0 a a a 0 0 0 a 0 0 0 0 U
Stony Off-Road 1973 a a a a a a 0 a a 0 0 0 a a 0 a 0 a 0 a
Sable Road 1973 a a a 0 5 0.242 2 0.097 a a a a a 0 0 0 26 1.256 2 0.097
N
N
W
r ''''\rrr:7J ,;,...,r:-l rJ C"J CTJJ r::J [TJ .-'"l c-~::J C'"'J c--J Ll rJ 1:---:1 I~n c-Jj~L
Table A7.Actual numbers and minilmJlIl densities of Dall sheep observed durin9 each pl,ot observation period.total day.Mount McKinley National Park.Alaska.
PLOT OBSERVATION PERIOD (TOTAL UAY)
2 3 4 5
Animals Groups Animals Groups AniNls Groups Anima 1s Groups Anir..als GrOUjjS
Density Density Density Density Density liens i ty Dens ity liens i ty Dens ity Density
No.(Per knh No.(Per knh No.(Per km2 )No.(Per knh No.(Per ki)No.(Per k(2 )110.(Per knll)No.(Per km l )No.(Per kml )No.(Per k(2 )
19100 Road 1973 77 5.033 10 0.654 11 0.719 4 0.261 30 1.961 4 0.261 32 2.092 3 0.196 14 0.915 0.065
Igloo Road 1974 9 0.558 5 0.327 27 1.765 4 0.261 19 1.242 3 0.196 17 1.111 2 0.131 15 0.980 2 0.131
19100 Off-Road 1973 80 4.167 4 0.208 112 5.833 10 0.521 94 4.896 11 0.573 72 3.750 5 0.260 102 5.312 4 0.208
Igloo Off-Road 1974 30 1.562 7 0.365 3 0.156 ,1 0.520 62 3.229 6 0.312 48 2.500 6 0.312 115 4.427 4 0.208
Highway Road 1973 8 0.447 2 0.112 9 0.503 0.056 3 0.168 0.056 0 0 0 0 0 0
()0
Hi9hway Off-Road 1973 0 0 0 0 14 1.082 2 0.155 0 0 0 0 0 0 0 (j 4'0.310 0.078
Stony Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Stony Off-Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ()
Sable Road 1973 2 0.338 0.145 5 0.242 2 0.097 0 0 0 0 0 0 0 0 26 1.256 2 0.097
N
N
~
r ['J c-i
''1,--,flu II ..---.,rl :~i~r-J c-llL:'!""J ("~C1 c-J CTl c-"l r-::J c=-J !
(
Table A2..Actual numbers and minimum densities of brown bears observed during each plot observation period.first-search.Mount McKinley National Park.Alaska.
PLOT OflSERV/,TIOfl PlRlOD (FIRST-StARCH)
2 3 4 ;,
Animals Groups Animals Groups Animals Groups Animals Groups Anin.a1s Groups
Density Density Density Density Dens I ty lJens i ty Density Oens I ty Densl ty IJensi ty
flo.(Per kr.h flo.(Per km2)flo.(Per km2)No.(Per km2)/lo.(Per kch 110.(Per knh No.(Per knh No.(Per km 2)flo.(Per km2)i/o.(Per l:ml )
Igloo Road 1973 1 0.065 0.0&5 0.065 0.065 0 0 0 0 0 0 0 0 0 0 0 0
Igloo Road 1974 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Igloo Off-Road 1973 0.052 0.052 0.052 0.052 3 0.156 2 0.104 0.052 0.052 0 0 0 0
Igloo Off-Road 1974 4 0.208 2 0.104 0 0 0 0 0 0 0 0 0.052 0.052 6 0.312 3 u.156
highway Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0.056 0.056 0 0 II 0
Highway Off-Road 1973 0 0 0 0 0 0 0 0 0.078 0.078 1 0.078 0.078 0.078 v.078
Stony Road 1973 0 0 0 0 0 0 0 0 3 0.242 0.081 0 0 0 0 0.081 u.Oa1
Stony Off-Road 1973 6 0.612 2 0.204 0 0 0 0 0 0 0 0 0 0 0 0 0 0
(J U
sable Road 1973 3 0.145 0.048 10 0.483 4 0.193 6 0.290 4 0.193 3 0.145 0.047 0 0 0
(i
N
Nc....,
r :----,l.)
Table 1.9.ACtual numbers and ~inimum densities of brown bears observed during each plot observation period.total day.
Mount McKinley National Park.Alaska.
PLOT OBSERVATIOli PtRIOU (TOTAL UAY)
2 3 4 )
I.nimals Groups Animals Groups Animals Groups Animals Groups Animals Gro"ps
iiensity Density Uens i ty Density Uensity Uensity Uensity Llellsity uensity Dens i ty
No.(Per ki)No.(Per kJn2)No.(Per kli)No.(Per kJn2)/lo.(Per knh No.(Per ki)110.(Pe~knh lio.(Per kJn2)110.(Per krn2 )Ho.(Fer ~.})
Igloo Road 1973 0.065 0.065 0.065 0.065 0 0 0 0 0 0 0 0 0 0 0 u
Igloo Road 1974 0 0 0 0 0 0 0 0 4 0.261 0.0&5 0 0
0 0 0 0 U U
1:;1,;,;:-:ff-Road 1973 4 0.2OS 2 0.104 0.052 0.052 3 0.156 2 0.104 3 0.1)6 2 0.104 0 0 0 0
Igloo Off-Road 1974 4 0.208 2 0.104 0 0 0 0 0 0 0 0 0.052 0.05'6 0.312 3 u.156
Hi9hway Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0.0)6 0.056 0 0 0 u
Highway Off-Road 1973 0 0 0 0 0 0 0 0 0.078 0.078 0.078 0.078 0.078 0.078
Stony Road 1973 0 0 0 0 0 0 0 0 3 0.242 O.OUl 0 0
0 0 0.01l1 u.OBl
Stony Off-Road 1973 6 0.612 4 0.204 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0
Sable Road 1973 7 0.338 3 0.145 11 0.531 5 0.242 6 0.290 4 0.193 6 0.290 2 0.097 6 0.290 2 0.097
rl r'u ru rr--,~J ",_;.J ~i.:"U
Table Al0,Actual numbers and minimum densities of red foxes observed during each plot observations period,first-search,
Mount McKinley lIatillna1 Park,Alaska.
PLOT OBSERVATIOll PERICJli (FIRST-S!.AKCH)
2 3 4 5
Anir.131s Groups Animals Groups Animals Groups Animals Groups Animals Groulis
Uensity Uensity Uensity Density Density liens i ty Density LJensity liens i ty uensity
(Per kli)No.(Per km2 )No.(Per ki)No.(Per kr:h No.(Per km2 )No.(Per ki)110.2 (Per km2 ).lo.(Per k.,,2),10.(Per'Kr.h110.(Per kill )No.
Igloo Road 1973 2 0.131 2 0.131 3 0.196 0.065 0 0 0 0 0 0 0 0 0 0 0 II
Igloo Road 1974 0 0 0 0 5 0.327 2 0.131 0 0 0 0 0.065 0.06:)0 0 0 0
Igloo Off-Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Igloo Off-Road 1974 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Highway Road 1973 0 0 0 0 0 0 0 0 0.056 0.056 2 0.112 0.050 0 0 U 0
Highway Off-Road 1973 0 0 0 0 0 0 0 0 0.078 0.078 0 0 0 0 0 0 0 0
Stony Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 O·0 '0 IJ U
Stony Off-Road 1973 0.102 0.102 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Sable Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 IJ
N
N
'-l
r:r:J r .......1'----'l ,j
.....--,
l J
Table All.Actual numbers and minimum densities of red foxes observed during each plot observation period.total day,
!'!ountMcKinley flational Park,Alaska.
PLOT OBSERVATIOfl PERIOU (TOTAL liAY)
2 3 4 5
Animals Groups Anin:a 1s Groups Animals Groups Anin:als Groups Animals urou~s
Oensity Density liensity uensity Dens ity liensity liens ity liens i ty iJensity Liens ity
flo.(Per krh flo.(Per kC12 )No.(Per km2)No.(Per km2)flo.(Per km2 )No.(Per km2)flo.(Per km2)No.(Per kch flo.(Per km2)Iio.(Per m£)
Igloo Road 1973 2 0.131 2 0.131 3 0.195 0.065 0 0 0 0 0 0 0 0 0 0 0
0
Igloo Road 1974 0 0 0 0 5 0.327 2 0.131 0 0 0 0 0.065 0.065 0 0 0 0
Igloo Off-Road 1973 0.052 0.052 0 0 0 0 0 0 0 0 0
0 0 0 0 0 u 0
Igloo Off-Road 1974 0 0 0 0 a 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0
Highway Road 1973 0.056 0.056 0 0 0 0 0.056 0.056 2 0.112 0.056 0 0
0 il
Highway Off-Road 1973 0 0 0 0 0 0 0 0
0.078 0.078 0 0 0 0 0 0
0 0
Stony Road 1973 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 U U
Stony Off-Road 1973 0.102 0.102 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
sable Road 1973 0 0 0 0 0 0 0 0 0.O4!l 0.048 0 0 0 0 0 0 0 U
rj
Table A12.Total number of each species of large maQ~21 observed during each shuttle-tour round-trip from Riley Campground
to Eielson Visitor Center.Mount NcKinley National Park.Alaska.
DATE TYPE BUS CARIBOU NOOSE OALL SHEEP BROWil BEAR REO FOX I\OLF
Number Number Number Number Number Number Number Number Number Number Number /lumber
Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups Anir:lals Groups
1973
May 26 0600 Shuttl e 18 9 4 4 30 7 5 2 1 1 0 0
27 1500 Shuttle 18 3 12 10 53 .13 7 3 0 0 0 0
28 1500 Shuttle 3 3 12 6 18 5 3 1 4 2 0 0
June 3 1500 Shuttle 47 7 10 8 16 6 4 2 l:l 1 0 0
8 1500 Shuttle 24 6 13 9 48 7 12 4 1 1 0 0
9 1500 Shuttle 47 7 13 8 54 10 2 2 12 5 1 1
10 0600 Shuttle 118 12 11 8 67 11 3 3 5 2 0 0
11 1500 Shuttle 85 13 5 4 87 10 14 6 5 3 1 1
12 1500 Shuttle 146 17 13 g 12 3 9 5 5 2 1 1
17 1500 Shuttle 25 11 12 7 16 5 11 5 2 2 0 0
19 0700 Shuttle 12 7 7 7 49 8 2 2 4 3 0 0
23 0400 Tour 3 2 9 6 32 6 12 4 0 0 0 0
24 0';00 Tour 7 2 2 1 59 3 11 4 7 1 0 0
27 0400 Tour 2 2 4 2 43 6 13 5 4 2 0 0
29 1500 Shu ttl e 10 5 1 1 39 3 4 2 2 2 0 0
July 1 1500 Sh~ttle 16 8 2 1 19 2 4 1 0 0 0 0
3 0400 Tour 6 1 1 1 5 1 3 1 0 0 1 1
4 0700 Shuttle 12 3 6 2 32 3 1 1 1 1 0
o·
10 1500 Shuttle 4 2 1 1 24 5 11 5 0 0 0 0
12 0600 Shuttle 10 4 1 1 60 4 11 5 6 2 0 0
13 0400 Tour 18 8 4 3 35 3 5 3 4 2 0 0
17 0400 Tour 14 5 12 5 34 2 5 3 2 2 0 0
19 0600 Shuttle 10 2 3 2 30 5 13 5 1 1 0 0
24 0700 Shuttle 15 10 0 0 24 6 6 4 0 0 0 0
26 0400 Tour 21 6 9 5 6 1 5 3 1 1 1 1
27 0700 Shuttle 26 5 2 1 11 1 8 4 1 1 0 0
31 0400 Tour 9 5 5 4 22 3 4 2 2 1 0 0
Aug.1 0700 Shuttle 10 5 2 2 36 4 0 0 2 2 0 0
2 1500 Shuttle 13 5 0 0 20 2 5 3 0 0 0 0
5 1500 Shuttle 12 3 6 4 16 4 10 4 3 2 0 0
7 0700 Shuttle 11 5 0 0 75 8 9 3 0 0 0 0
8 0400 Tour 11 6 5 4 17 1 3 1 1 1 0 0
r "';"V ,./'I..,j,C:J r -4"""1'1 rn L:"J c:r=J CJ]r=J CTJ r-J r~::J r-J r-J rl rJ l'""":J r-J rl r-JJ
Table A12 (continued)
DATE TYPE BUS CARIBOU NOOSE VALL SHEEP BROWN BEAR RED FOX WOLF
Number Number Number Number Number Number I~umber Number Number Humber ilumber IlUl:lber
Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups
1973 (cont.)
AU9.16 1500 Shuttle 0 0 16 8 2 1 11 5 1 1 0 0
17 0700 Shuttle 3 3 0 0 36 3 13 5 0 0 0 0
18 0600 Shuttle 5 2 4 2 22 3 7 3 0 0 0 0
23 0400 Tour 7 5 9 4 70 11 9 3 2 2 0 0
24 1500 Shuttle 7 7 2 2 30 3 9 3 0 0 a 0
25 0700 Shutt1 e 3 2 5 3 62 6 6 2 0 ·0 0 a
31 0700 Shuttle 0 0 8 4 46 5 6 2 0 0 0 0
Sept.2 0700 Shuttle 0 0 4 2 28 6 3 1 0 0 0 0
3 0700 Shuttle 0 0 3 2 18 3 0 0 0 0 0 0
1974
May 26 0400 Tour 36 6 11 5 29 9 16 6 2 2 0 0
31 1500 Shuttle 9 6 10 6 8 3 1 1 0 0 0 0
June 5 0400 Tour 3 1 0 0 4 2 0 0 0 0 0 0
8 1500 Shuttle 0 0 4 3 26 4 7 3 0 0 0 0
9 0400 Tour 3 1 6.6 34 6 6 2 0 0 0 0
14 1500 Shuttle 4 2 4 2 36 8 10 5 1 1 0 0
19 0400 Tour 12 2 6 4 46 3 0 0 1 1 0 0
26 1500 Shuttle 10 10 8 6 45 3 9 5 1 1 0 0
28 0400 Tour 11 5 2 2 60 6 9 3 1 1 0 0
29 1500 Shuttle 12 6 2 1 77 6 5 3 2 2 0 0
3D 1500 Shuttle 4 1 1 1 59 4 3 1 0 0 0 0
July 6 0400 Tour 32 6 10 7 34 4 8 4 2 2 0 0
18 1500 Shuttle 10 9 5 4 40 5 10 5 0 0 0 0
23 0400 Tour 20 6 11 3 23 3 4 2 0 0 0 0
27 1500 Shuttle 8 8 1 1 41 6 3 1 0 0 0 0
30 0400 Tour 23 5 5 4 4 2 12 4 0 0 0 0
31 0400 Tour 34 8 3 2 57 6 9 3 1 1 0 0
Aug.3 1500 Shuttle 25 11 5 3 37 5 5 3 0 0 0 0
5 0400 Tour 11 8 8 4 51 5 0 0 0 0 0 0
9 1500 Shuttle 7 4 7 5 22 2 0 0 0 0 0 0
16 0400 Tour 25 7 8 6 19 2 3 1 0 0 0 0
17 1500 Shuttle 22 3 1 1 37 2 0 0 0 0 0 0
18 0400 Tour 16 4 25 16 33 4 0 0 0 0 0 0
19 1500 Shuttle 11 3 7
4 4 2 3 1 0 0 0 0
25 0400 Tour 4 2 8 6 30 2 3 1 0 0 0 0
30 0400 Tour 0 0 8 5 26 2 3 1 0 0 0 0
31 1500 Shuttle 0 0 1 1 69 5 3 1 1 1 0 0
Sept.1 0400 Tour 17 8 7 6 57 10 0 0 1 1 0 0
2 1500 Shuttle 2 2 0 0 10 2 4 2 0 0 0 0
N
W
0
--:-J
Table A13.Reacti ons of car;bout observed at vary;ng di stances from the road dur;ng shutt1 e-tour tri ps.
to the bus and visitors i numbers of animals and groups.1973-1974.Mount McKinley National
Park.Alaska.
DISTANCE REACTION OF CARIBOU TO BUS AND VISITORS TOTAL
FROM
ROAD UNKNOWN NONE MILD STRONG
(m)
%Grand %Grand
Number Number Number NUll'ber Number Number Number Number Number Number Total Total
Ani ma 1s Groups Animals Groups Animals Groups Animals Groups Ani ma 1s Groups Ani ma 1s Groups
0-25 4 3 4 3
12 5 21 11 42 21 4 6
0-100 26 10 84 41 42 24 90 33 243 104 21 30
0 2 1 0 0 11 4 13 8 26 11 2 3
1-25 2 2 4 3 1 1 9 5 16 11 1 3
26-50 4 1 18 11 14 6 24 8 60 26 5 8
51-75 2 1 16 7 12 9 3 2 33 18 3 5
76-100 16 5 46 20 4 4 42 12 108 40 9 12
101-200 9 7 151 55 47 9 28 8 235 7~20 23
201-300 3 3 140 36 16 6 9 3
]68 47 14 14
301-400 32 4 87 24 12 2 6 2 137 32 12 9
401-500 3 2 113 19 8 2 0 0 124 23 11 7
501-750 6 2 119 23 0 0 0 0 125 25 11 7
751-1000 24 5 30 11 0 0 1 1 55 17 5 5
>1000 0 0 Y2 16 0 0 0 0 92 16 8 5
TOTAL 103 33 816 225 125 43 135 48 1179 342
Note:Some groups are counted under more than one reaction class if various animals in the group
exhibited different types of reactions.Thus.the number of groups entered under the
different reaction classes may total more than the actual total number of groups given.
The percentages of animals and groups exhibiting each type of reaction appear in Table 6.
,...---..,
•j
N
W.......
rl w:-;~\c-TI Cl tI':J L",1.1 ~,J)Ll r ~.----,r-J [-.i .----..,r--J ;'J ,.---,:--J r-JII,,,.J -'
Table A14.Reactions of caribou.observed during shuttle-tour trips.to various actions of the bus and
vi s itors;numbers of animals and groups.1973-1974.Mount McKinley National Park.Alaska.
ACTION OF REACTION OF CARIBOU TOTAL
BUS AND
VISITORS UNKNOHN NONE t~ILD STRONG
;~Grand ,;Grand
Number Nunber Number Number Number Nu:mer Number Number Number Number Total Total
Ani ma 1s Groups Animals Groups Animals Groups Animals Groups Ani rna 1s Groups Animals Groups
Pass >15 mph 14 6 148 44 12 6 6 180 57 15 17
Pass <15 mph 28 3 26 7 4 2 59 12 5 4
StoP.people 42 21 513 146 79 26 77 29 711 219 60 64
remain on
bus,quiet
Stop,people 2 13 5 9 3 15 6 39 14 3 4
remain on
bus,noisy
StoP.people 4 93 17 18 5 23 7 138 30 12 9
off bus on
road,quiet
StoP.people 0 0 5 2 0 0 6 3 <1
off bus on
road,noi sy
Stop,people 13 13 3 3 9 38 6 3 2
walk off
road,quiet
Stop.people 0 0 5 0 0 3 8 2
<1 <1
walk off
road,noisy
TOTAL 103 33 816 225 125 43 135 48 1179 342
Note:Some groups are counted under more than one reaction class if various animals in the group
exhibited different types of reactions.Thus.the number of groups entered under the
different reaction classes may total more than the actual total number of groups given.
Nw
N
Lt..'
,
C"""l .,...--....,
!C""]I "~
Table A15.Reactions of caribou,observed during shuttle-tour trips,to various actions of
the bus ~nd visitors;percent of animals and groups.1973-1974,Mount McKinley
National Park.Alaska.
ACTION OF BUS REACTION OF CARIBOU TOTAL
AND VISITORS KNOWN
NONE MILD STRONG REACTIONS
Percent Percent Percent Percent Percent Percent Number Number
Animals Groups Animals Groups Animals Groups Animals Groups
Pass >15 mph 89 86 7 12 4 2 166 51
Pass <15 mph 84 70 13 20 3 10 31 10
StoP.people 77 73 12 13 12 14 669 200
remain on
bus;qui et
Stop,people 35 36 24 21 41 43 37 14
remai n on
bus,noisy
Stop,people 69 59 13 17 17 24 134 29
off bus on
road ,qui et
Stop,people 83 67 0 0 17 33 .6 3
off bus on
road,noisy
Stop,people 52 60 12 20 36 20 25 5
~Ial k off
road,qui et
Stop,people 62 50 a a 38 50 8 2
walk off
road,noi sy
TOTAL 76 72 12 14 12 14 1076 313
Note:"Groups"may total over 100%.since some groups were counted under more than one
reaction class when various animals in the group reacted differently.
N
W
W
~t,.I ."j
,--..,.j
Table A16.Reactions of caribou,observed within 200 m of the road during shuttle-tour trips,to various
actions of the bus and visitors;number of animals and groups,1973-1974.Mount McKinley
National Park.Alaska.
REACTION OF CARIBOUACTIONOF
BUS AND
VISITORS UNKNOHN NONE 11ILD STRONG
TOTAL
%Grand %Grand
Number Number Number Number Number Number Number Number Number Ilumber Total Tota 1
Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups
Stop,people 24
remain on .
bus.quiet
Stop,people 2
remai n on
bus.noi sy
StoP.people 4
off bus on
road.qui et
Stop,people 0
off bus on
rO'ad.noisy
StoP.people 0
walk off
road,quiet
Stop,people 0
walk off
road.noisy
Pass >15 mph
Pass <15 mph
3
2
2
12
o
o
o
20
12
157
13
28
5
o
o
12
3
66
5
8
2
o
o
7
4
63
9
3
o
3
o
4
2
20
3
3
o
o
6
70
15
14
9
3
27
6
4
36
19
314
39
49
6
12
3
19
6
122
14
15
3
2
8
4
66
8
10
2
<1
10
3
67
8
8
2
<1
TOTAL 35 17 235 96 89 33 .119 42 478 182
Note:Some groups are counted under more than one reacti on cl ass if vari ous animal sin the group
exhibited different types of reactions.Thus,the number of groups entered under the
different reaction classes may total more than the actual total number of groups given.
The percentages of animals and groups exhibiting each type of reaction appear in Table 8.
(--
l I)rj
N
W
(J"l
rl r---'/lJ,
Table Ala.Reactions of moose.observed during shuttle-tour trips.to various actions of the bus and
visitors;number of animals and groups.1973-1974.Mount McKinley National Park.•Alaska.
ACTION OF REACTIOn OF ~<l()OSE TOTAL
BUS AND
VISITORS UNKNmm rWNE MILD STRONG
%Grand %Grand
Number Nur:ber Number Nur.ter Nurrber Number Number Number Number Number Total Total
Animals Groups Animals Groups Animals Gro ups An i ma 1s Groups Animals Groups Animals Groups
Pass >15 mph a 6 29 28 2 2 5 4 44 40 11 15
Pass <15 mph 1 9 6 3 3 14 11 3 4
StoP.people 44 26 148 92 46 27 47 34 285 174 69 65
remain on
bus.qui et
Stop,people 2 3 2 6 6 14 9 25 18 6 7
remain on
bus.noi sy
StoP.people 6 5 11 7 a 4 16 9 41 22 10 8
off bus on
road,qui et
Stop,people 0 0 3 2 1
off bus on
road,noi sy
Stop,people 0 0 0 0 0 0 0 0 0 a 0 0
walk off
road,qui et
Stop,people a a 0 0 a 0 a 0 0 0 0 0
walk off
road,noi sy
TOTAL 62 40 201 136 64 41 8,5 59 412 267
Note:Some groups are counted under more than one reaction class if various animals in the group
exhibited different types of reactions.Thus,the number of·groups entered under the
different reaction classes may total more than the actual total number of groups given.
N
W
O"l
r1 ,,
:-\~"l\;~
\...-_...-../
/
i~
Table A19.Reactions of moose,observed during shuttle-tour trips,to various actions
of the bus and visitors;percent of animals and groups,1973-1974,Mount
McKinley National Park,Alaska.
ACTION OF REACTION OF MOOSE TOTAL
BUS AND Kr;o\1N
VISITORS NONE MILD STRONG REACTIONS
Percent Percent Percent Percent Percent Percent Number Number
Animals Groups Animals Groups Ani rna 1s Groups Animals Groups
Pass >15 mph 81 82 6 6 14 12 36 34
Pass <15 mph 69 60 8 10 23 30 13 10
Stop,people 61 61 19 18 20 23 241 150
remain on
bus,quiet
Stop,people 13 12 26 35 61 53 23 17
rema;n on
bus,noisy
Stop,people 31 35 23 20 46 45 35 20
off bus on
road,qui et
Stop,people 50 50 50 50 0 0 2 2
off bus on
road,noi sy
Stop,people 0 0 0 0 0 0 0 0
walk off
road,qUi et
Stop,people 0 0 0 0 0 0 0 0
1'1 alk off
road,noisy
TOTAL 57 58 18 18 24 25 350 233
Note:"Groups"may total over 100%since some groups were counted under more than one
reaction class when various animals in the group reacted differently.
L:....J ~<_J ',-"---1(_~l
.~
:---J
Table A20.Reactions of moose,observed within 200 m of the road during'shuttle-tour trips,to various
actions of the bus and visitors;number of animals and groups,1973-1974,Mount McKinley
National Park,Alaska.
ACTIOtI OF REACTION OF MOOSE TOTAL
BUS AND
VISITORS UNKNQ\oJN NQrJE t~ILD STRONG
%Grand %Grand
Number Number Number Number Number Nurrber Number Number Number
Number Total Total
Animals Groups Animals Groups Ani ma 1s Groups Ani ma 1s Groups Animals Groups Animals Groups
Pass >15 mph 7 5 16 15 2 2 5 4 30 26 9 12
Pass <15 mph 5 4 3 3 10 9 3 4
Stop,people 35 21 104 68 39 24 43 32 221 140 68 69
remai n on
bus,quiet
Stop,people 2 5 5 14 9 22 16 7 7
remai n on
bus,noisy
Stop,people 6 5 11 7 8 4
16 9 41 22 12 10
off bus on
road,quiet
Stop,people 0 0 3 2
off bus on
road,noisy
Stop,people 0 0 0 0 0 0 0 0 0
0 0 0
\~alk off
road,quiet
Stop,people 0 0 0 0 0 0 0 0 0
0 0 0
walk off
road,noisy
TOTAL 52 34 138 96 55 37 81 57 327 215
Note:.Some groups are counted under more than one reaction class if various animals in the group
exhibited different types of reactions.Thus,the number of groups ente'red under the
different reacti on cl asses may total more than the actual number of groups gi ven.The
percentages of animals and groups exhibiting each type of reaction appear in Table 20.
N
W
00
,
-1(,
'iii..,'
C,j r--:tlj.w ..".,w
,/
C"'J (J --"\.,J
Table A21.Reactions of Da11 sheep,observed at varying distances from the road during shuttle-tour
trips,to the bus and visitors;number of animals and groups,1973-1974,Mount McKinley
National Park.Alaska.
DISTANCE REACTION OF DALL SHEEP TO BUS AND VISITORS TOTAL
FROr~
ROAD UNKNOWN NONE MILD STRONG
(m)
%Gra nd %Grand
Number NUr.1ber Number Number Number Number Number Number Number Number Total Total
Animals Groups Animals Groups Animals Groups Ani ma 1s Groups Ani rna 1s Groups Animals Groups
0-25 0 0 10 6 9 4 32 8 51 16 2 5
0-100 4 2 51 15 24 9 40 11 120 34 5 10
0 0 0 0 0 3 1 21 3 24 4 1
1
1-25 0 0 10 6 6 3 11 5 27 12 1 4
26-50 0 0 5 3 4 2 7 2
16 7 1 2
51-75 3 1 24 3 2 1 0 0 29 5 1 2
76-100 1 1 12 3 9 2 2 1 24 7 1
2
101-200 19 4 14 4 0 0 1 1 34 9 1 3
201-300 8 2 105 18 2 1 8 2 123 23 5 7
301-400 8 2 185 21 3 2 1 1
197 25 8 8
401-500 38 4 283 36 28 3 0 0 349 43 14 13
501-750 7 3 233 38 3 1 0 0 243 42 io 13
751-100 22 4 888 88 0 0 0 0 910 92 37 28
>1000 5 1 446 54 0 0 0 0 451 55 18 17
TOTAL 111 22 2205 274 60 16 51 15 2427 324
Note:Some groups are counted under more than one reaction class if various animals in the group
exhibited different types of reactions.Thus.the number of groups entered under the
different reaction classes may total more than the actual total number of groups given.
The percentages of animals and groups exhibiting each type of reaction appear in Table 28.
~!J~..-..~, 1
Table A22.Reactions of Oall sheep,observed during shuttle-tour trips,to various actions of the bus
and visitors;number of animals and groups,1973-1974,Mount McKinley National Park,Alaska.
ACTION OF REACTION OF DALL SHEEP TOTAL
BUS AND
VISITORS UNKNOWN NONE MILD STRONG
%Grand %Grand
Number Number tlumber Number Number Number Number Number Number NUClber Total Total
Animals Groups Animals Groups Animals Groups Ani ma 1s Groups Animals Groups AniJr.als Groups
Pass >15 mph 44 9 486 76 6 3 0 0
536 88 22 27
Pass <15 mph a a 71 8 5 3 0 a 76 11 3 3
Stop,people 34 10 1174 140 41 7 20 8 1269 163 52 50
remain on
bus,quiet
Stop,people 0 0 0 0 20 2 21 3
remain on
bus,noisy
Stop,people 19 2 453 46 8 3 9 4 489 53 20 16
off bus on
road,quiet
Stop,people 14 16 3 0 0 0 '0 30 4
off bus on
road,noisy
Stop,people 0 0 4 0 0 2 6 2 <1 <1
walk off
road,quiet
Stop,people 0 0 0 0 0 0 0 0 0
0 0 0
I-Ia 1k off
road,noi sy
TOTAL 111 22 2205 274 60 16 51 15 2427 324
Note:Some groups are counted under more than one reaction class if various animals in the group
exhibited different types of reactions.Thus,the nUJrber of groups entered under the
different reaction classes may total more than the actual total number of groups given.
II ".....
Ii
,
k.J {'.~
....,J
Table A23.Reactions of Da11 sheep,observed during shuttle-tour trips.to various actions
of the bus and visitors;percent of animals and groups,1973-1974,Mount
McKinley National Park,Alaska.
ACTION OF REACTION OF DALL SHEEP TOTAL
BUS AND KIWI-iN
VISITORS NONE NILD STRONG REACTIONS
Percent Percent Percent Percent Percent Percent Nu~ber Number
Animals Groups Animals Groups Animals Groups Anil'1als Groups
Pass >15 mph 99 86 14 0 0 492 79
Pass <15 mph 93 73 7 27 0 0 76 11
StoP.people 95 91 3 5 2 5 1235 154
remai n on
bus.qui et
StoP.people 5 33 0 0 95 67 21 3
remai n on
bus.noi sy
Stop,people 96 90 2 6 2 8 470 51
off bus on
road,quiet
Stop,people 100 100 0 0 0 0 16 3
off bus on
road,noisy
StoP.people 67 50 0 0 33 50 6 2
~Ialk off
road,qui et
Stop,people 0 0 0 0 0
0 0 0
walk off
road,noi sy
TOTAL 95 91 2 5 2 5 2316 303
Note:"Groups"may tota 1 over 100%,since some groups were counted under more than one
reaction class when various animals in the group reacted differently.
f,-.-1I....
Table A24.Reactions of Dall sheep,observed within 200 m of the road during shuttle-tour trips,to
various actions o~the bus and visitors;numbers of animals and groups,1973-1974,Mount
McKinley National Park,Alaska.
ACTION OF REACTION OF DALL SHEEP TOTAL
BUS AND
VISITORS UNKNOl-lN Nm:E't·HLD STRONG
%Grand %Grand
Number Number Nur.:ber Number Number Number Number Number Number Number Tcta1 Total
Animals Groups Animals Groups Ani ma 15 Groups Animals Groups Ani ma 1s Groups Animals Groups
Pass >15 mph 9 2 5 2 2 0 0 16 5 10 12
Pass <15 mph 0 0 2 5 3 0 0 7 4 4 9
Stop,people 14 4 46 10 15 4 14 7 89 23 58 53
remain on
bus,quiet
Stop,people 0 0 0 0 0 0 20 2 20 2 13 5
rema in on
bus,noisy
Stop,people 0 0 9 5 2 8 3 19 8 12 19
off bus on
road,qui et
Stop,people 0 0 3 0 0 0 0 3 2 2
off bus on
road,noi sy
Stop,people 0 0 0 0 0 0 0 0 0 0 0 0
walk off
road,quiet
Stop,people 0 0 0 0 0 0 0 0 0 0 0 0
walk off
road,noi sy
I
TOTAL 23 6 65 19 24 9 42 12 154 43
Note:Some groups are counted under more than one reaction class if various animals in the group
exhibited different types of reactions.Thus,the number of groups entered under the
different reaction classes may total more than the actual total number of groups given.
The percentages of animals and groups exhibiting each type of reaction appear in Table 30.
N
..f::>
N
r-j \IT:;,(-,'r-'"J ['~.J t;r""J tr:'1 r-"rt:J r -C:-J C-:J ..---~~'1 [ )
,~r--J t-"1,"","),j ',{,J l I,,'.,.,
Table A25.Reactions of brown bears,observed at varying distances from the road during shuttle-tour
trips,to the bus and visitors;numbers of animals and groups,1973-1974,Mount McKinley
National Park,Alaska.
DISTANCE REACTION OF BRO\1N BEARS TO BUS Arm VISITORS TOTAL
FROM
ROAD UNKNm1N NONE MILD STRONG
(m)
%Grand %Grand
Number Number Number Number Nurrber NUr.lber tlumber Number Number Number Total Total
Animals Groups Animals Groups Ani ma 1s Groups An i ma 1s Groups Animals Groups Animals Groups
0-25 3 1 0 0 5 5 16 6 24 12 6 7
0-100 9 3 48 20 15 11 39 14 111 46 27 25
0 0 0 0 a 2 2 13 5 15 7 4 4
1-25 3 1 0 0 3 3 3 1 9 5 2 3
26-50 0 a 7 3 5 3 3 1 15 7 4 4
51-75 a 0 23 9 2 2 6 2 31 11 7 6
76-100 6 2 18 8 3 1 14 5 41 16 10 9
101-200 8 5 50 20 4 2 11 5 73 32 18 18
201-300 10 4 56 25 5 2 0 0 71 31 17 17
301-400 0 0
31 15 3 1 3 1 37 17 9 9
401-500 6 2 25 13 0 0 0 0 31 15 7 8
501-750 0 0 33 15 0 0 0 0 33 15 8 8
751-1000 3 1 31 11 0 0 3 1 37 13 9 7
>1000 3 1 19 11 0 0 a 0 22 12 5 7
TOTAL 39 16 293 130 27 16 56 21 415 180
Note:Some groups are counted under more than one reaction class if various animals in the group
exhibited different types of reactions.Thus,the number of groups entered under the
di fferent reacti on c1 as ses may total rrore than the actual total number of groups gi ven.
The percentages of animals and groups exhibiting each type of reaction appear in Table 39.
I~
\J
Table A26.Reactions of brown bears,observed during shuttle-tour trips,to various actions of the bus
and visitors;numbers of animals and groups,1973-1974,r~ount McKinley National P'ark,Alaska.
ACTION OF REACTIOtJ OF BROWN BEARS TOTAL
BUS Nm
VISITORS UNKNO,IN NONE MILD STRONG
%Grand %Grand
Number Number NUlT.ber Number Number Number Number Number Number Number Total Total
Animals Groups Animals Groups Animals Groups Animals Groups Animal s Groups Animals Groups
Pass >15 mph 3 14 8 0 0 0 0 17 9 4 5
Pass <15 mph 0 0 2 2 a 0 a a 2 2 <1
Stop,people 29 12 180 80 16 9 32 13 257 113 62 63
remai n on
bus.quiet
Stop,people 22 8 2
2 9 3 34 14 8 8
remai n on
bus.noi sy
Stop,people 3 72 31 9 5 12 4 96 39 ;23 22
off bus on
road,quiet
Stop,people 3 3 a 0 3 9 ':3 2 2
off bus on
road,noisy
Stop,people a a a a 0 a a a a a a a
walk off
road,quiet
Stop,people a a a a a 0 0 0 0 0 0 0
walk off
road,noi sy
TOTAL 39 16 293 130 27 16 56 21 415 180
Note:Some groups are counted under more than one reaction class if various animals in the group
exhibited different types of reactions.Thus,the nu~ber of groups entered under the
different reaction classes may total more than the actual total number of groups given.
N
-1:>0
-1:>0
~"'I r-t ~r:--:;L':"7J r-:->,.,.......-..,,..-.LC1 r j "'-~r--'!l (J r-:\----1 rJ .,..-..;...,c-1 .~l J J[,;:I _)C \,I _~~'"."I 4;.....,...l .,j J j I ,..,}
Table A27.Reactions of bro~m bears,ohserved during shuttle~tour trips,to various actions
of the bus and visitors;percent of animals and groups,1973-1974,Mount
McKinley National Park,Alaska.
ACTION OF REACTIO~1 OF BROInl BEARS TOTAL
BUS AND KrW~:N
VISITORS NOtlE MILD STRO:1G P.!:ACTW!S
Percent Percent Percent Percent Percent Percent r'lu:nber :Jumber
Animals Groups Animals Groups Animals Groups Animals Groups
Pass >15 mph 100 100 0 0 a 0 14 8
Pass <15 mph 100 100 0 0 0 0 2 2
Stop,people 79 78 7 9
14 13 228 102
remai n on
bus,qui et
Stop,people 67 62 6 15 27 23 33 13
remai n on
bus,noi sy
Stop,people 81 89 10 14 13 11 89 35
off bus on
road,quiet
Stop,people 50 50 0 0 50 50 6 2
off bus on
road,noisy
StoP.people 0 0 0 0 0 0 0 0
I','alk off
road,quiet
Stop,people 0 0 0 0 0 0 0 0
~Ialk off
road,noisy
TOTAL 79 80 7 10.15 13 372 162
Note:"Groups"may total over 100%,since some groups ~Iere counted under more than one
reacti on cl ass when va ri ous animals in the gro,up reacted differently.
N
.j:::o
U1
Table A28.Reactions of brown bears,observed within 200 m of the road during shuttle-tour trips,to
various actions of the bus and visitors;nurrbers of animals and groups,1973-1974,Mount
McKinley National Park,Alaska.
REACnDrl OF BROWN BEARSACTIONOF
BUS AND
VISITORS WlKNOWN NONE ~1ILD STRONG
TOTAL
%Grand %Grand
Number Number Number Number Number Number Number Number Number Number Tota 1 Tota 1
Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups Animals Groups
Pass >15 mph
Pass <15 mph
Stop,people
remain on
bus,qui et
Step,people
remai n on
bus,noi sy
Stop,people
off bus on
road,qui et
Stop,people
off bus on
road,noi sy
Stop,people
walk off
road,quiet
Stop,people
~Ialk off
road,noi sy
TOTAL
3
o
10
o
3
o
o
17
o
4
o
o
o
7
o
53
9
32
3
o
o
98
o
21
3
14
o
o
40
o
o
11
2
6
o
o
o
19
o
o
7
2
4
o
o
o
13
o
o
26
9
12
3
o
o
50
o
o
11
3
4
o
o
19
4
o
100
21
50
9
o
o
184
2
o
43
9
20
3
o
o
77
2
o
54
11
27
5
o
o
3
o
56
12
26
4
o
o .
Note:Sorre groups are counted under more than one reaction class if various animals in the group
exhi bited different types of reacti ons.Thus,the number of groups entered under the
different reaction classes may total more than the actual total number of groups given.The
percentages of animals and groups exhibiting each type of reaction appear in Table 40.
'--.j
[
rto
[
Table A29.Research truck road census trips,dates of trips and
condition of road access,Mount McKinley National
Park,Alaska.
247
CENSUS TRIPS 1973 CENSUS TRIPS 1974
Sept.3 II
Sept.4 II
Sept.7 /I
Sept.8 /I
l]
C
C
TJ
[
f'
Date
May 21
May 21
May 22
May 23
May 24
May 24
June 21
Augus t 30
Augus t 31
Sept.1
Sept.2
Sept.3
Sept.4
Sept.8
Sept.9
Sept.22
Sept.23
Road Access
Road closed to public
beyond Mi 1e 30
/I
/I
II
/I
II
Road open to public
buses and private
vehicles wi~h permits
II
II
Road open to all
private vehicles
wi thout permi ts
II
II
II
II
II
II
/I
Date
May 25
May 29
r~ay 30
May 30
June 1
August 28
August 29
August 31
Sept.1
Road Access
Road closed to public
beyond Mi 1e 30 .
Road open to puhlic
and private vehicles
wi th pe rmi ts
II
/I
/I
/I
/I
Road open to all
private vehicles
wi thout permi ts
/I
Jtr -
i:'",'ii,'I ./
r---1
\ j
Table A30 Average daily traffic,traveling east plus west combined,through Savage River Guard Station 1974,Mount McKinley National Park,Alaska.
Week
begin-
ning
Time of day (hour)
1800 --2400
Nps Publ1c Prlvate
veh.bus veh.Total
Total.
NPS Public Priv~te
veh.bus veh.Total
June 2 0 2.0 1.5
June 9 0.6 3.4 3.4
June 16 0.4 2.7 2.7
June 23 0.3 4.0 3.7
June 30 0.9 5.0 4.0
July 7 0.7 5.4 1.9
3.5 6.5 20.5 11.0 37.0 5.0 10.6 17.4 33.0 2.3 3.0 9.0 14.3 14.8 36.1 38.9 89.8
7.4 4.7 14.3 19.4 38.4 6.0 11.0 23.6 40.6 4.0 6.4 18.0 28.4 15.3"34.4 64.4 114.1
5.8 6.6 19.3 24.8 SO.7 5.2 16.9 32.5 54.6 3.8 6.9 20.7 3J.4 16.0 45.8 80.7 142.5
8.0 5.5 20.6 27.6 53.7 5.5 20.0 35.4 60.9 4.3 7.7'21.7 33.7 15.6 52.3 88.4 156.3
9.9 5.4 22.5 27.7 55.6 6.0 19.6 37.3 62.9 2.4 10.2 20.4 33.0 15.0 57.3 99.4 171.7
8.0 5.5 22.0 23.5 51.0 7.1 22.1 32.0 61.2 4.6 9.5 21.5 35.6 17.9 59.0 78.9 lj5.8
July 14 0.7 5.8 4.2'10.7 6.3 25.1 25.7 57.1 6.3 22.1 36.7 65.1 3.5 13.0 26.2 42.7 16.8 66.0 92.8 175.6
July 21 0.6 5.8 2.7
July 28 0.4 5.3 2.3
August 4 0.3 5.6 1.2
August 11 0.7 6.0 2.4
August 18 0.4 5.1 3.2
August 25 0.3 3.8 2.1
Sept.1*0 4.0 1.5
9.1 6.2 25.1 22.8 54.1 7.2 22.0 37.7 66.9 4.1 12.2 21.3 37.6 18.1 65.1 84.5 167.7
8.0 3.1 26.4 30.3 59.8 4.3 21.8 35.7 61.8 3.3 11.7 26.0 41.0 11.1 65.2 94.3 170.6
7.1 4.7 24.6 24.1 53.4 5.4 22.6 39.6 67.6 4.9 7.6 23.7 36.2 15.3 60.4 88.6 164.3
9.1 3.8 26.9 26.9 57.4 4.8 20.4 33.6 58.8 5.3 10.1 19.6 35.0 14.6 ·63.4 82.5 160.5
8.7 4.9 20.4 35.2 50.6 4.7 19.3 37.9 61.9 4.6 9.7 18.1 32.4 14.6 54.5 84.5 153.6
6.2 3.3 20.6 21.8 45.7 4.4 18.8 36.0 59.2 4.2 8.6 22.0 34.8 12.2 51.8 82.9 146.9
5.5 2.0 19.5 38.5 60.0 5.0 16.0 45.5 66.5 3.5 12.5 27.0 43.0 10.5 52.0 132.5 195.0
*Data for only two days.
[
[
[
['
[
,['~
.,'.-...J
[
[
F
r
G
L
LITERATURE CITED
Ables,E.D.1975.Ecology of the red fox in North America.Pages
216-236 in M.W.Fox,ed.The wild canids.Van Nostrand Reinhold
Co.,N.Y.
Allison,L.N.1971.Activity and behavior of red foxes in Central
Alaska.M.S.Thesis.Univ.of Toronto,Toronto,Ontario.92 pp.
Altmann,11'1.1958a.The flight distance in free-ranging big game.
J.Wildl.Nanage.22(2):207-209.
•1958b.Social integration of the moose calf.Anim.Behav.
-----r4(3,4):155-159.
Andersen,R.1971.Effects of human disturbance on Da 11 sheep.Alas ka
Coop.Wildl.Research Unit Quart.~Rept.22(3):23-27 •
Banfield,A.~.J.F.1954.Prel iminary investtgations of the barren-
ground caribou,Part II:Life history,ecology,and utilization.
Can.Wildl.Servo Wildl.Manage.Bull.Sere 1,No.lOB.112 pp.
__..,....1974.The relationship of caribou migration lJehavior to pipe-
line construction.Pages 797-804 in V.Geist and F.Walther,eds.
The behavior of ungulates and its relation to management.Vol.2.
Int.Union Conserve Nat.Hat.Resourc.Publications flew Series
No.24,Morges,Switzerland.
Barnes,V.G.,Jr.and O.E.Bray.1967.Final report:Population
characteristics and activities of black bears in Yellowstone
National Park.Submitted to the U.S.Nat.Park Servo Colorado
Coop.Wildl.Research Unit,Colorad~State Univ.,Fort Collins.
199 pp.
Baskin,L.fil.1974.Nanagement of ungulate herds in relation to
domestication.Pages 530-541 in V.Geist and F.Walther,eds.
The behavior of ungulates and its relation to management.Vol.2.
Int.Union Conserve Nat.Nat.Resourc.Publications Hew Series
No.24,Morges,Switzerland.
Bergerud,A.T.1974.The role of environment in the aggregation,
movement,and disturbance behavior of caribou.Pages 552-584
in V.Geist and F.Walther,eds.The behavior of ungulates and
its relation to management.Vol.2.Int.Union Conserve Nat.
Nat.Resourc.Publications New Series No.24,Morges,Switzerland.
249
.....,
j.
250
Buskirk 1 S.1975.U.S.Dept.of the Interior,National Park Service,
Case incident record do.1514.Unpubl.Ufficial Files,Hount
NcKinley National Park,U.S.(Jat.Park Servo ~pp.
Calef,G.W.and G.M.Lortie.1973.Observations on the Porcupine
caribou herd.Pages 1-127 in Interim report no.3 towards an
environmental impact assessment of the portion of the MacKenzie
gas pipeline from Alaska to Alberta.Environmental Protection
Board,Winnipeg,t~nitoba •
Carbaugh,B.,J.P.Vaughan,E.D.Bellis,and II.B.Graves.197b.
Distribution and activity of \'/hite-tailed deer along an interstate
highway.J.Wildl.Manage.39(3):570~81.
Carbyn,L.N.1974a.Wolf population fluctuations in Jasper National
Park,Alberta,Canada.Biol.Conserve 6(2):94-101.
_______•1974b.Wolf predation and behavioral interactions with elk
and other ungulates in an area of high prey density.Unpubl.Can.
Wildl.Servo Prepared for Parks Canada.Jasper National Park.
233 pp.
Cauble,E.C.1975.Chronology of grizzly bear attacks on~humans from
Aug.,1961 through r·lay,1975,r·lt.NcKinley National Park.Unpubl.
Official Files,Nount tkKinley I~ational Park,U.S.Hat.Park
Servo 3 pp.
Chapman,R.C.(in prep.).The effects of human disturbance on wolves
(Canis 'lupus L.).N.S.Thesis.Univ.of Alaska,Fairbanks •
.Child,K.N.1973.The reactions of barren-ground caribou (Rangifer
tarandus granti)to simulated pipel ine and pipel ine crossing
structures at Prudhoe Bay,Alaska.A Completion Report to the
Alaska Coop.~Jildl.Research Unit,Univ.of Alaska,Fairbanks.
49 pp.
and P.C.Lent.1973.The reaction of reindeer to pipeline
-----simulation at Penny River,Alaska.Interim Report,Alaska Coop.
Wildl.Research Unft,Univ.of Alaska,Fairbanks.30 pp.
Chrisler,L.1956.Observations of wolves hunting caribou.J.
Mamn~l.37(3):337-346.
Cole,G.F.1972.Preservation and management of grizzly bears in
Yello~stone National Park.Pages 274-288 in S.Herrero,ed.
Bears --their biology and management.Int.Union Conserve I~at.
Nat.Resourc.Publ ications New Series I'Jo.23,Norges,Switzerland.
Couey,F.H.1950.Rocky f10untain bighorn sheep of t10ntana.Fed.Aid
in Hildl.Rest.Proj.·l-R,Bull.2,f10ntana Fish and Game Commis-
sion.90 pp.
Cowan,I.M.1974.Management implications of behavior in the large
herbivorous mammals.Pages 921-934 in V.Geist and F.Walthers,
eds.The behavior of ungulates and its relation to management.
Vol.2.Int.Union Conserve Nat. Nat.Resourc.Publications New
Series No.24,Morges,Switzerland.
Cowan,1.not D.G.Chapman,R.S.Hoffman,D.R.r~cCullough,G.fl.
Swanson,and R.B.Weeden.1974.Report of the committee on the
Ye11o\'/stone grizzlies.National Academy of Sciences,Division of
Bio10nical Sciences,Assembly of Life Sciences,National Research
Counci 1.61 pp.
Craighead,F.C.,Jr.and J. J.Craighead;1966.Trailing Yellow-
stone's grizzlies by radio.Natl.Geogr.~·lag.134(8):252-267.
Craighead,F.C.,Jr.and J.J.Craighead.1972a.Data on grizzly
bear denning activities and behavior obtained by using wildlife
telemetry ..Pages 84-106 in S.Herrero,ed.Bears --their
bi 01 ogy and management.Int.Union Conserv.~la t.Nat.Resourc.
Publications New Series No.23,Morges,Switzerland.
______~.1972b.Grizzly bear prehibernation and denning activities as
determined by radio tracking.Hildl.t-1ongr.No.32.35 pp.
Craighead,J. J.and F.C.Craighead,Jr.1972.Grizzly bear-man
rel ati onshi ps in Yellowstone tlati ona 1 Park.Pages 304-332 in
S.Herrero,ed.Bears --their biology and management.Int.
Union Conserve Nat.Nat.Resourc.Publications r·le\'/Series No.23,
Morges,Switzerland.
Curatolo,J.fl.1975.Factors influencing local movements and beha-
vior of barren-ground caribou (Rangifer tarandus g2?anti).N.S.
Thesis.Univ.of Alaska,Fairbanks.146 pp.
and G.D.Moore.1975.Home range and population dynamics of
-------grizzly bears (UrsU8 arotos L.)in the Eastern Brooks Range,
Alaska.Pages 1-79 in R.D.Jakimchuk,ed.Studies of large
mammals along the proposed ~'lacKenzie valley gas pipeline route
from Alaska to British Columbia.Arctic Gas Biol.Rept.Sere
Vol.32.
Dar1inq,F. F.1937.A herd of red deer.Oxford Univ.Press,London.
215 pp.
251
..,
252
Dean,F.C.(in press).Aspects of grizzly bear ecology in Mount
McKinley National Park.Paper given at the Third International
Conference on Bear Research and Nanagement held Hay 31-June 1,
1974 in Binghamton,New York.
De Bock,E.A.and D.Surrendi.1974.Some aspects of the ecology of
the Porcupine caribou herd.Preliminary report prepared for the
Task Force on Northern Oil Development, Ottawa.n.p.(Cited in
Geist 1975,original not seen.)
Denniston,R.H.1956.Ecology,behavior and population dynamics of
the Wyoming or Rocky ~buntain moose.Zoologica 41:105-118.
de Vos,A.1958.Summer observations on moose behavior in Ontario.
J.Manma1.39(1):128-139.
___•1960.
grounds.
Behavior of barren ground caribou on their calving
J.Wi1d1.Manage.24:250-258.
j
Dixon,J.S.1938.Birds and marrmals of ('-10unt McKinley National Park,
Alaska.Fauna of the National Parks of the U.S.,Faun&Ser.No.
3.U.S.Govt.Printing Office,Wash.,D.C.236 pp.
Dorrance,H.J.,P.J.Savage,and D.E.Huff.1975.Effects of sno\'1-
mobiles on \'/hite-tailed deer.J.Wildl.Hanage.39(3):563-69.
Downing,C.1975.Hildlife activity at the HcKin1ey Park area garbage
dUlilps.Unpubl.report submitted to the u.S.Nat.Park Servo and
Mount HcKinley National Park from the Alaska Coop.Park Studies
Unit,Univ.of Alaska,Fairbanks.67 pp.
Dugmore,A.R.1913.The romance of the Newfoundland caribou.J.B.
Lipincott Co.,Phil.191 pp.
Erickson,A.W.1965.The brown-grizzly bear in Alaska,its ecology
and management.Fed.Aid in Wi1dl.Rest.Proj.Rept.5.Alaska
Dept.Fish and Game.42 pp.
Ericson,C.A.1972.Some preliminary observations on the acoustic
behavior of semi-domestic reindeer (Rangifer tarandus tarandus)
with emphasis on intraspecific cOri1llunication and the mother-calf
relationship.M.S.Thesis.Univ.of Alaska,Fairbanks.121 pp.
Espmark,Y.1972.Behavior reactions of reindeer exposed to sonic
booms.Deer.Jour.Brit.Soc.2(7):800-802.
...,
--"
Faro,J.and S.Eide.1974.t·1ana~eJl1ent of ~1ctleil River State Game
Sanctuary for nonconsurnptive use of Alaska brown bears.Paqes
113-118 in Proc.Fifty-fourth Ann.Conf.Western Ass.State Game
and Fis.h Comm.,Albequerque,N.t·t,July 16-19,1974.
Freuchen,P.and F.Salomonsen.1958.The arctic year.G.P.Put-
nam's Sons,N.Y.438 pp.
Geist,V.1963.On the behavior of North American moose (AZees aZees
andersoni Peterson 1950)in British Columbia.Behav.20(3-4):377-
416.
253
.J
•1971a.A behavioral approach to the management of wild ungu-
---=-lates.Pages 413-424 in E.Duffey and A.S.Hatt,eds.The sci-
entific management of animal and plant communities for conserva-
tion.Brit.Ecol.Soc.Symp.No.11.Blackwell Sci.Publ.,
Oxford.
1971b.Is big game harassment harmful?Oilweek 22(17):12-13.
·1971c.t·1ountain sheep:a study in behavior and evolution.-----~Univ.of Chicago Press,Chicago,Ill.383 pp.
•1975.Harassment of large mammals and birds --with ·a cri-
---:-ti que of the research submi tted by Arcti c Gas Study Ltd.on the
subject.Unpubl.report to the Berger Commission.69 pp.
Haber,G.C.1968.The social structure and behavior of an Alaskan
wolf population.M.S.Thesis.Northern Michigan Univ.,Mar-
quette.197 pp.
•1972a.The McKinley Project:behavioral-ecological studies
---of wolves and prey in a subarctic biosystem.Unpubl.Univ.of
British Columbia,Vancouver.21 pp.
__-::-.1972b.t10unt McKinley Park and north boundary vrildlife prob-
lems --a proposal.·Unpubl.Univ.of British Columbia,Vancou-
ver.21 pp •
Harper,F.1955.The barren ground caribou of Keewatin.Univ.of
Kansas Press,Lawrence.163 pp.
Heimer,W.,A.C.Smith,S.Linderman,and P.D.Shaughnessy.1972.
Dall sheep movements and mineral lick use.Proj.Rept.Fed.Aid
Wild.Rest.Proj.H-17-3 and U-17-4.Pages 1-17 in L.Nichols
and W.Heimer.Sheep Report.Vol.13.Alaska ·nept.Fish and
Game,Juneau..
-,
.J
j
-------....._.---_..-----_._-------_..--_._----------------
254
Hernning,J.E.1971.The distribution and movement patterns of cari-
bou in Alaska.Alaska Dept.Fish and Game Tech.Bull.No.1.
60 pp.
Henshaw,J.1970.Consequences of travel in the rutting of reindeer
and caribou (Rangifel'tarandus).Anim.Behav.18(2):256-2~8.
Herrero,S.1970.Human injury infl icted by grizzly bears.Science
170(3958):593-598.
__,.....1972.Aspects of evolution and adaptation in American black
bears (Ul'SUS americanU$Pallus)and brown and grizzly bears (u.
al'ctos Linn~)of North funerica.Pages 221-231 in S.Herrero,ed.
Bears --their biology and management.Int.Union Conserve Ilat.
Nat.Resourc.Publications New Series No.23,Borges,S\'litzerland.
Houston,D.G.1968.The Shiras moose in Jackson Hoie,Hyoming.
Tech.Bull.No.1,Grand Teton Natural History Assoc.in coop.
with the U.S.Nat.Park Servo 110 pp.
•1974.Aspects of social organization of moose.Pages 690-696---:-in V.Geist and F.Walther,eds.The behavior of ungulates and
its relation to management.Vol.2.Int.Union Conserve lJat.
Nat.Resourc.Publ ications New Series No.24,norges,Switzerland.
Jakimchuk,R.D.,E.A.De Bock,H. H.Russell,and G.P.Semenchuk.
1974.A study of the Porcupine caribou herd,1971.11~pp.in
R.D.Jakimchuk,ed.The Porcupine caribou herd --Canada.
Arctic Gas Bio1.Rept.Sere Vol.4•
Johnson,A.S.1972.Man,grizzly and national parks.Natl.Parks
Conserve Mag.46(2):10-15.
Jones,F.F.,R.F.Batchelor,H.R.r·lerriam,and L.A.Viereck.1963.
Sheep and goat investigations.Vol.III.Ann.Proj.Segment
Rept.Fed.Aid in Wildl.Rest.Act.Proj.W-6-R-3,Work Plan E.
Alaska Dept.Fish and Game,Juneau.47 pp.
Kelsall,J.P.1957.Continued barren-ground caribou studies.Wildl.
Manage.Bull.Sere 1,No.12.Can.Uept.Northern Affairs and
National Resources.148 pp.
__~.1968.The migratory barren-ground caribou of Canada.Can.
Wildl.Servo Honogr.No.~.Queen's Printer,Ottawa.340 pp.
Klein,U.R.1971.Reaction of reindeer to obstructions and distur-
bances.Science 173(3995):393-398.
255
Klein,D.R.1974.The reaction of son~northern n2nma1s to aircraft
disturbances.Pages 377-3H3 in Proc.XI Int.Congr.Game Biolog-
ists,Stockholm,Sweden,Sept.3-7,1973.
Knowlton,F.F.1960.Food habitats,movements and populations of
moose in the Gravelly Nountain,Nontana.J.Wild1.Hanage.
24(2):162-170.
Krott,P.and G.Krott.1963.Zum verha1ten des Braunbaren (Ursus
arctos L.1758)in den Alpen [in German,English summary].z.f.
Tierpsycho1.20(2):160-206.
Lenarz,M.1974.The reaction of Da1l sheep to an FH-1l00 helicopter.
Ch.3,14 pp.in R.D.Jakimchuk,ed.The reaction of some malilnla 1s
to ai rcraft and compressor station noi se di sturbance.Arctic Gas
Bio1.Rept.Sere Vol.23.
Lent,P.C.1964.Calving and related social behavior in the barren
ground caribou.Ph.D.Thesis.Univ.of Alberta,Edmonton.
220 pp.
Lentfer,J.W.,J.R.Blum,S.H.Eide,and L.H.Miller.1966.Bear
studies.Fed.-Aid in Wi1dl.Rest.Rept.Proj.W-6-R-6.Alaska
Dept.Fish and Game,Juneau.54 pp.
LeResche,R.E.1966.Behavior and calf survival in Alaska nIDose.
H.S.Thesis.Univ.of Alaska,Fairbanks.85 pp.
•1974.Noose migrations in North America.Natural iste Can.---~10·1:393-415.
Linderman,S.1972.A report on the sheep study at the Dietrich River
headquarters.Appendix 3.Proj.Rept.Fed.Aid Wild1.Rest.
Proj.W-17-3,W-17-4.13 pp.in L.Nichols and W.Heimer.Sheep
Repurt,Vol.13.Alaska Dept.Fish and Game,Juneau.
•1974.Ground tracking of arctic grizzly bears.Final Rept.,
------~Fed.Aid in Wildl.Rest.Proj.W-17-6,Job 4.12 R.Alaska Dept.
Fish and Game,Juneau.24 pp..
Lloyd,H.G.
Fox,ed.
1975.The red fox in Britain.Pages 207-215 in M.W.
The wild canids.Van Nostrand Reinhold Co.,N.Y.
r·1acPherson,A.H.1965.The barren-ground grizzly and its survi va 1 in
Canada.Can.Audobon 27(1):2-8.
n
256
Nartinka,C.J.1974a.Population characteristics of grizzly bears
in Glacier National Park,Hontana.J.r··'1ammal.55(1):21-29.
______~.1~74b.Preserving the natural status of grizzlies in Glacier
Hationa1 Park.Wildl.Society Bull.2(1):13-17.
fvlcCann,J.L.1956.Ecology of the mountain sheep.Am.Nidl.Nat.
o 56(2):297-3~5.
McCourt,K.H.,J.D.Feist J D.Doll,and J. J.Russell.1974.Dis-
turbance studies of caribou and other.mammals in the Yukon and
Alaska,1972.Arctic Gas Biol.Rept.Sere Vol.5.246 pp.
McCourt,K.H.and L.P.Horstman.1974.The reaction of barren-
ground caribou to aircraft.Ch.1,36 pp.in R.U.Jakimchuk,
ed.The reaction of some mammals to aircraft and compressor
station noise disturbance.Arctic Gas Biol.Rept.Sere Vol.23.
McCullough,D.R.1969.The Tule elk:its history,behavior,and
ecology.Univ.of Cal.Press,Berkeley.209 pp..~
NcKnight,D.,ed.1975.Annual report of survey-inventory activities,
Part I:Noose,deer and elk.Vol.5,1970-1975.Fed.Aid in
14ildl.Rest.Alaska Dept.Fish and Game,Juneau.n.p.·
Nc~1illan,J.F.1954.Some observations on moose in Yello\'/stone Park.
Am.Hidl.Nat.52(2):392-399.
Mech,L.D.1970.The wolf:the ecology and behavior of an endangered
species.Nat.Iiist.Press,Garden City,N.Y.384 pp.
Nemphis State University (for the Environmental Protection Agency).
·1971.Effects of noise on wildlife and other animals.U.S.Govt.
Printing Office,I·lash.,U.C.74 pp.
Miller,F.L.,C.J.Jonkel,and G.D.Tessier.1972.Group cohesion
and 1eadershi p response by barren-ground cari bou to man-made bar-
riers.Arctic 25(3):193-201.
Mundy,K.R.D.and D.R.Flook.1973.Background for managing grizzly
bears in the national parks of Canada.Can.Wildl.Servo kept.
Sere No.22.Information Canada.Uttawa.35 pp.
Nurie,A.1934.The moose of Isle Royale.Univ.f1ich.Nus.Zool.f<1isc.
Pub1.25.44 pp.
~,,
,,j
j
257
Murie,A.1944.The wolves of Mount McKinley.Fauna of the National
Parks of the U.S.,Fauna Sere No.5.U.S.Govt.Printing Office,
Wash.,D.C.238 pp •
•1961.A naturalist in Alaska.The Devin-Adair Co.,N.Y.---'::-30~2 pp.
__~.1962.Nanmmls of [>lount r'lcKinley Hational Park,Alaska.Haunt
NcKinley Natural History Ass.in coop.\vith the Pisani Printing
Co.,U.S.Nat.Park Serv.,San Francisco.56 pp.
___•1963.Birds of r·lount NcKinley,Alaska.Hount t'!cKinley Natu-
ral History Ass.in coop.with the U.S.Nat.Park Servo Pisani
Printing Co.,San Francisco.86 pp.
f.1urie,O.J.1935.Alaska-Yukon caribou.N.Amer.Fauna Sere Ho.54.
U.S.Govt.Printing Office,Wash.,U.C.93 pp.
Nurphy,E.C.1973.,An age structure and a reevaluation of the popu-
1a ti on dynami cs of Ua 11 sheep (OVis daZU daZZi).N.S.•Thes is.
Univ.of Alaska,Fairbanks.113 pp.
Pearson,A.M.1975.The northern interior grizzly bear Ursus arctos
L.Can.Wildl.Servo Rept.Sere No.34.Information Canada,
Ottawa.86 pp.
Peters,R.P.1974.Wolf-sign:scents and space in a wide-ranging
predator.Ph.D.Thesis.Univ.of Michigan,Ann Arbor.228 pp.
Peterson,R.L.1955.The North American moose.Univ.of Toronto
Press,Toronto,Ontario.280 pp.
Peterson,R.O.and D.L.Allen.1974.Ecological studies of the wolf
on Isle Royale.Unpubl.Fourth Ann.Rept.,1973-1974,to Nat.
Park Servo 14 pp.
Pienaar,U.De V.1968.The ecological significance of roads in a
national park.Koedoe 11:169-187.
Price,R.1972.Effects of human disturbance on Dall sheep.Alaska
Coop.Wildl.Research Unit.Quart.Rept.22(3):23-28.
Pruitt,W.O.,Jr.1960.Behavior of the barren-ground caribou.Univ.
of Alaska Biol.Pap./Jo.3.44 pp.
•1965.A flight releaser in wolf-caribou relations.J.-~Nanmal.46(2):350-351.
258
Quimby,R.1974.Grizzly bear.Ch.2,97 pp.in Nammal studies in
northeastern Alaska with emphasis within the Canning River drain-
age.Can.Arctic Gas Biol.Rept.Sere Vol.24.
Rausch,R.A.1959.Son~aspects of population dynamics of the rail-
belt moose populations,Alaska.M.S.Thesis.Univ.of Alaska,
Fairbanks.81 pp.
Reed,D.F.,T.N.Woodard,and T.M.Pojar.1975.Behavioral re-
sponse of mule deer to a highway underpass.J.Wildl.t'-Ianage.
39(2):361-367.
Reynolds,H.V.1974.North Slope grizzly studies.Vol.1.Proj.
Rept.,Fed.Aid in Wildl.Rest.Proj.W-17-6,Jobs 4.8R,4~9R,
4.l0R,and 4.l1R.Alaska Dept.Fish and Game.27 pp.
Reynolds,P.C.1974.The effects of simulated compressor station
sounds on Dall sheep using mineral licks on the Brooks Range,
Alaska.Ch.2,82 pp.in R.D,.Jakimchuk,ed.The reaction of
some mammals to aircraft and compressor station noise disturbance.
Arctic Gas Biol.Rept.Sere Vol.23.~
Riegelhuth,R.1966.Grizzly bears and human visitation.M.S.
Thesis.Colorado State Univ.,Fort Collins.80 pp.
Rue,L.L.,III.1969.The world of the red fox.J.B.Lippincott
Co.,N.Y.204 pp.
Russell,A.1967.Grizzly country.Knopf,N.Y.302 pp.
Schoonmaker,H.J.1968.The world of the grizzly bear.J.B..Lippin-
cott Co.,N.Y.190 pp.
Sheldon,C.1930.The wilderness of Denali.Scribner1s,N.Y.412 pp.
Skoog,R.O.1963.Caribou report.Fed.Aid'in Wildl.Rest.Proj.
W-6-R-4.Alaska Dept.Fish and Game,Juneau.31 pp.
•1968.Ecology of the caribou (Hangifer tarandus granti)in------A~laska.Ph.U.Thesis.Univ.of Cal.,Berkeley.699 pp.
Smith,D.R.1954.The bighorn sheep in Idaho --its status,life
history,and management.Fed.Aid Wildl.Rest.Idaho Proj.99-R.
Idaho Uept.Fish and Gan~Hildl.Bull.Ho.1.Boise,Idaho.
154 pp.
Sokal,R.R.and F.J.Rohlf.1969.Biometry.'WI H.Freeman and Co.,
San Francisco.776 pp.
l
J
1
I
I
-J
I
j
!
-.J
1
j
j
----.-------------,
259
Storm,G.L.1972.Population dynamics of red foxes in Horth Central
United States.Ph.D.Thesis.Univ.of r·linnesota,Ninneapolis-
St.Paul.227 pp.
Stringham,S.F.1971.Calf behavior and the cO\/-calf bond in Illoose.
Alaska Coop.Hildl.Research Unit Quart.Rept.22(4):12-32.
•1974.Nother-infant relations i~semi-captive Alaskan moose
--"7"'{AZaes aZaes gigas).H.S.Thesis.Univ.of Alaska,Fa·irbanks.
140 pp.
Symington,F.1965.Tuktu --the caribou of the northern mainland.
Can.Wildl.Serv.,Ottawa.92 pp.
Thomson,B.R.1972.Reindeer disturbance.Deer.Jour.Brit.Deer
Soc.2(8):882-883.
1973.Hild reindeer activity,Hardangervidda,1971.Report
from the grazing project of the Norwegian IBP comrilittee.Statens
viltunders~kelser,Direktoratet for Jakt,Viltstell og Ferskvanns-
fiske,Trondheim,Norway.76 pp.~.
Troyer,W.A.1974.Hinter moose census,Nount HcKinley Hational
Park;1974.Unpubl.Official Files,Nat.Park Serv.,Anchorage,
Alaska.4 pp.
Troyer,H.A.and R.J.Hensel.1964.Structure and distribution of
a Kodiak bear population.J.Wild1-f·lanage.28(4):769-772.
Tyrrell,J.W.1898.Across the sub-Arctic of Canada,a journey of
3,200 miles by canoe and snowshoe through the barren lands.-
T.Fisher Unwin,London.280 pp.
U.S.Dept.of Corrmerce.1970.Climatic summaries of resort areas --
Nount HcKinley Park,Alaska.U.S.D.C.Publication No.21-49-1-
5 pp.
U.S.Dept.of Interior,Alaska Planning Group.1974.Final impact
statement:proposed Nt.r·1CKinley National Park additions,Alaska.
-------687 -pp.-- -- - - - - - - - - - - - - ---- - ---- -----. - - - - - - - - - - - - - - - - - - - -.----- - - -
Valkenburg,P.1976.A study of the brown bear (Ursus aratos)in the
proposed northeastern addi tion to Ivlount NcKi nl ey National Park.
M.S.Thesis.Univ.of Alaska,Fairbanks.88 pp.
Van Wormer,J.1972.The world of the moose.J.B.Lippincott Co.,
N.Y.160 pp.
-...,
!
1
260
Videsott,R.1967.Impact eco10gique du tourisme et des 10isirs dans
1es parcs nationaux ita1iens [in French,English summary].Pages
118-120 in Towards a new relationship of man and nature in temper-
ate lands.Part I.Ecological impact of recreation and tourism
upon temperate environments.Int.Union Conserve Hat.Hat.
Resourc.,Norges,S\'Iitzer1and.
Viereck,L.A.and E.L.Little,Jr.1972.Alaska trees and shrubs.
Agriculture Handbook No.410.Forest Service,U.S.Dept.of Agri.
U.S.Govt.Printing Office,Wash.,D.C.265 pp.
von Uexkull,J.1957.A stroll through the \'lorld of animals and men.
Pages 5-80 in Schiller,C.H.,ed.Instinctive behavior.Inter-
national Universities Press,Inc.,N.Y.
Ward,A.L.,J.J.Cupa1.,A.L.Lea,C.A.Oakley,and R.W.Weeks.
1973.Elk behavior in relation to cattle grazing,forest recrea-
tion,and traffic.Pages 327-337 in J.B.Trefethen,ed.Trans.
38th N.Amer.Wi1d1.and Nat.Res.Conf.,Wi1dl.Manage.Inst.,
Wash.,D.C.
Washburn,B.1971.A tourist guide to Hount NcKin1ey.Alaska North-
west Pub 1.Co •.,Anchorage,Al aska.79 pp.
Welles,R.E.and F.B.~Jelles.1961.The bighorn of Death Valley.
Fauna of the National Parks of the U.S.,Fauna Sere Ho.6.U.S.·'.
Govt.Printing Office,Wash.,O.C.242 pp.
White,R.G.,B.R.Thomson,T.Skog1and,S.J.Person,D.E.Russell,
D.F.Holleman,and J.R.Luick.1975.Ecology of caribou at
Prudhoe Bay,Alaska.Pages 151-201 in J.Brown,ea.Ecological
investigations of the tundra biome in the Prudhoe Bay region,
Alaska.Univ.of Alaska Biol.Pap.,Spec.Rept.'Jo.·2.
VJhitten,K.R.1975.Habitat relationships and population dynamics
of Dall sheep (avis daUi daUi)in t1t.NcKinley r-Jational Park,
Alaska.N.S.Thesis.Univ.of Alaska,Fairbanks.177 pp.
Hi1coxon,F.and R.Wilcox.1964.Some rapid approximate statistical
_procedures.Leder1e Laboratories,Pearl River,N.Y.60 pp.
Zhigunov,P.S.,ed.1961.Reindeer husbandry.Trans1.from Russian.
Israel Program for Sci.Trans1.,Jerusalem,1968.348 pp.