(DRAFT) - Taxonomy
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
TAXONOMY
NAME - CARIBOU, WOODLAND
OTHER COMMON NAMES - CARIBOU, WOODLAND;CARIBOU, MOUNTAIN;CARIBOU, MOUNTAIN, SELKIRK;CARIBOU, MOUNTAIN, WESTERN;CARIBOU and SELKIRK;CARIBOU;TSUUGWI;REINDEER
ELEMENT CODE -
CATEGORY - Mammals
PHYLUM AND SUBPHYLUM - CHORDATA,
CLASS AND SUBCLASS - MAMMALIA,
ORDER AND SUBORDER - ARTIODACTYLA,
FAMILY AND SUBFAMILY - CERVIDAE,
GENUS AND SUBGENUS - RANGIFER,
SPECIES AND SSP - TARANDUS, CARIBOU
SCIENTIFIC NAME - RANGIFER TARANDUS CARIBOU
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Woodland Caribou
Rangifer tarandus caribou (Banfield, 1961)
KINGDOM: Animal GROUP: Mammal
PHYLUM: Chordata CLASS: Mammalia
ORDER: Artiodactyla FAMILY: Cervidae
As members of the deer family, caribou in general (and in this
case woodland caribou) are even-toed ruminants distinguishable from
other members of the deer family primarily by foot adaptations
suitable for travel in deep snow, antler structure, the presence of
antlers on females, and coloration. The feet are large, and dew claw
width and length contributes greatly to the foot size to aid travel in
soft snow. Caribou are medium sized deer with bulls weighing up to
270 kg and cows to 135 kg (01). Recorded total lengths of bulls are
210 to 225 cm and cows are 190 to 205 cm. In bulls, heights at the
shoulder have been measured between 122 and 125 cm while cows measured
5 to 10 cm less (01). Antlers of male woodland caribou are large,
often palmate and many pointed. A palmate brow tine or shovel is
often present as well as a bez tine which extends from the main beam
forward just above the brow tine. Antler heights of 90 cm and widths
of 100 cm are normal. Female caribou antlers are much smaller and are
Taxonomy - 1 (DRAFT) - Taxonomy
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
often 20 to 40 cm long with 2 or 3 small tines. In winter caribou are
deep brown with greyish white on the belly, buttocks, and underside of
the tail, and on the neck of the male. The summer coat is dark
greyish-black in both males and females.
Seton (02) identified what has commonly been called mountain
caribou (Rangifer tarandus montanus Seton) in 1899. Allen (03)
identified the Osborn caribou (Rangifer tarandus osborni Allen) in
1902. In 1961 Banfield (04) revised the reindeer and caribou taxonomy
in Canada and the United States and the two subspecies along with
others were combined into the single subspecies Rangifer tarandus
caribou. The formerly recognized subspecies, R. t. montanus and R. t.
osborni, are the synonyms for the populations most closely associated
with the Federally listed population.
The following is a list of the taxonomic treatment of Rangifer
tarandus caribou (Gmelin) taken from Hall, 1981 (43):
1788 [Cervus tarandus] caribou Gmelin;
1829 Cervus tarandus var. sylvestris Richardson;
1847 Cervus hastalis Agassiz;
1884 Rangifer tarandus caribou True;
1896 Rangifer terraenovae Bangs;
1899 Rangifer montanus Thompson-Seton;
1901 Rangifer stonei J.A. Allen;
1902 Rangifer osborni J.A. Allen;
1912 Rangifer fortidens Hollister;
1914 Rangifer arcticus caboti G.M. Allen;
1915 Tarandus rangifer keewatinsis Millais;
1915 Tarandus rangifer labradorensis, Millais;
1915 Tarandus rangifer ogilvyensis Millais;
1919 Rangifer mcguirei Figgins; and
1935 Rangifer montanus selousi Barclay.
Specific concerning publications and type localities for the above
scientific names can be found in (43). The U.S. has listed the
woodland caribou (Rangifer caribou sylvestris) in the Convention on
Nature Protection and Wildlife Preservation in the Western Hemisphere
(1941).
Mountain caribou is the common name normally applied to the
caribou of southern British Columbia a local population of Rangifer
tarandus caribou, woodland caribou (05). Woodland caribou has been
used as the common name in the Federal Register and the "Checklist of
mammals of the United States and the U.S. Territories" (06). The
Shoshone indians referred to this caribou as "tsuugwi" which means
"reindeer" (42). Selkirk caribou or Selkirk mountain caribou are
other common names which are occasionally used (07). In Canada, this
subspecies is known as the western mountain caribou. There are local
names which refer to the remaining herd in the south Selkirk mountains
along the border between Washington, Idaho and British Columbia. It
is not considered a separate ecotype but merely identifies a local
population.
Type specimens of woodland caribou from Alberta, British
Columbia, Washington, Idaho and Montana may be found at the following
locations:
Taxonomy - 2 (DRAFT) - Taxonomy
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
Location No. of Specimens
----------------------------------------------------------------------
United States Natural Museum 18
American Museum of Natural History 15
University of British Columbia 9
British Columbia Provincial Museum 9
National Museum of Canada 6
University of Idaho 6
Carnegie Museum 5
University of Alberta 3
British Museum of Natural History 1
Nalurhistoriska Riksmuseet, Stockholm 1
This listing is taken from Banfield (04) and supplemented by a
check of museums in British Columbia, Washington, Idaho and Alberta.
It may be incomplete and the number of specimens may have changed in
some museums.
Taxonomy - 3 (DRAFT) - Status
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
STATUS
Coded Status
E: Federal Endangered
Commercial
Commercial/consumption
Game (Consumptive Recreational)
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The woodland caribou (Rangifer tarandus caribou) has been
designated an Endangered species pursuant to the Endangered Species
Act of 1973 (50 CFR 17.11; P.L. 93-205, 87 Stat. 884; 16 U.S.C.
1531-1540), as amended. The subspecies has this status in the States
of Idaho and Washington, and in that part of British Columbia (Canada)
bounded by the Canada-USA border, Columbia River, Kootenay River,
Kootenay Lake, and Kootenai River. Critical Habitat has not been
designated.
This subspecies is protected by the Lacey Act (P.L. 97-79, as
amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to import,
export, transport, sell, receive, acquire, or purchase any wild animal
(alive or dead including parts, products, eggs, or offspring):
(1) in interstate or foreign commerce if taken, possessed,
transported or sold in violation of any State law or
regulation, or foreign law; or
(2) if taken or possessed in violation of any U.S. law,
treaty, or regulation or in violation of Indian tribal law.
It is also unlawful to possess any wild animal (alive or dead
including parts, products, eggs, and offspring) within the U.S.
territorial or special maritime jurisdiction (as defined in
18 U.S.C. 7) that is taken, possessed, transported, or sold in
violation of any State law or regulation, foreign law, or Indian
tribal law.
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
USFS -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public lands under their control. Also responsible
for management/recovery on Forest Service lands. The
Forest Service is responsible for integrating
management, protection, and conservation of Federally
listed species into the Forest Planning process
Status - 1 (DRAFT) - Status
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
(36 CFR 219.19 and 219.20).
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
and to utilize their authorities to carry out programs for the
conservation of the species.
STATE STATUSES AND LAWS:
STATE: Idaho
DESIGNATED STATUS: Endangered
ADMINISTRATIVE AGENCY: Idaho Department of Fish and Game,
Idaho Fish and Game Commission.
STATE STATUTE: Idaho Code Sec. 36-103(a), 104(b), 201, and 1301;
STATE: Washington
DESIGNATED STATUS: Protected
ADMINISTRATIVE AGENCY: Washington Department of Game
STATE STATUTE: Title 77, Revised Code of Washington 77.16.050,
77.16.120 (in particular Game Code of Washington
77.12.320); WCA 232-12-470 and 480.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
The woodland caribou (R. tarandus sylvestris) was listed by the
U.S. in the Convention on Nature Protection and Wildlife Preservation
in the Western Hemisphere (1941). This would have presumably extended
coverage to the Federally listed population since R. t. sylvestris is
now included under R. t. caribou. However, the 1970 revision to the
Annex does not include this subspecies.
Canada lists R. t. caribou as Rare under the common name western
mountain caribou.
ECONOMIC STATUSES:
Prior to the establishment of indian reservations, Shoshoni
indians hunted the woodlnd caribou in the Sawtooth Mountains of
Central Idaho. The description of Northern Shoshoni utilization of
caribou as a subsistence resource is especially significant because it
expands the list of food resources exploited by these aboriginal
peoples (See R-Biology #42).
The caribou also once were taken at one time for sport and for
market hunting.
83/01/14:48 FR 01726/01726 - Emergency rule, listed in ID, WA, & B.C.
83/10/25:48 FR 49245/49249 - Proposed rule, listing as Endangered
84/02/29:49 FR 07390/07394 - Final rule, listed as Endangered
84/12/18:49 FR 49118/49119 - Review & Extend final rule to Montana
85/09/18:50 FR 37958/37967 - Review of vertebrate wildlife, MT popul.
87/01/21:52 FR 02239/02242 - Petit. findings & status rev., MT popul.
Status - 2 HABITAT ASSOCIATIONS
HABITAT - TERRESTRIAL
TERRESTRIAL
SOCIETY OF AMERICAN FORESTRY TYPES
SAF TYPE STAGE CLOSURE
Western white pine shrub--seedling
Western white pine young tree
Western white pine mature tree
Western white pine Old Growth
shrub--seedling
young tree
mature tree
Old Growth
Western larch shrub--seedling
Western larch young tree
Western larch mature tree
Western larch Old Growth
Lodgepole pine shrub--seedling
Lodgepole pine young tree
Lodgepole pine mature tree
Lodgepole pine Old Growth
Western white pine shrub--seedling
Western white pine young tree
Western white pine mature tree
Western white pine Old Growth
shrub--seedling
young tree
mature tree
Old Growth
Western larch shrub--seedling
Western larch young tree
Western larch mature tree
Western larch Old Growth
Lodgepole pine shrub--seedling
Lodgepole pine young tree
Lodgepole pine mature tree
Lodgepole pine Old Growth
LAND USE -
Transportation, communications, and Util
Cropland and Pasture
Mixed Rangeland
Evergreen Forest Land
Forested Wetland
Nonforested Wetland
Bare Exposed Rock
Tundra Land: Shrub Brush
Tundra Land: Herbaceous
Tundra Land: Bare Ground
Tundra Land: Wet
Habitat Associations - 1 LAND USE -
Tundra Land: Mixed
Snow or Ice: Perrenial Snowfields
NATIONAL WETLAND INVENTORY CODES
NWI NWICLS NWIMOD NWISPEC
Palustrine SS5
Palustrine SS4
Palustrine SS1
Palustrine FO5
Palustrine FO4
Palustrine EM5
Palustrine AB4
Palustrine AB2
Palustrine SS5
Palustrine SS4
Palustrine SS1
Palustrine FO5
Palustrine FO4
Palustrine EM5
Palustrine AB4
Palustrine AB2
COMMENTS ON HABITAT ASSOCIATIONS -
Woodland caribou inhabit a harsh mountain forest environment in
areas which typically receive 15 m of total snowfall per year.
Physical adaptations of caribou for walking on snow (21,22) permit a
seasonal habitat selection pattern which takes advantage of arboreal
lichens as a winter food source. Arboreal lichens can be found in
large quantities in subalpine forests and deep snow accumulations
raise caribou high into the tree canopy in winter allowing them to
take advantage of this seasonal food source (15,18,23,24). Great
seasonal differences in snow type and depth influence seasonal caribou
habitat selection and the literature identifies 5 seasonal habitat
selection patterns (15,17,18). These periods, early winter, late
winter, spring, calving, and summer/fall/rut are discussed
individually.
Early winter begins as snow begins to accumulate in the mountains
in mid-October. During this period the snow soon reaches depths of 50
to 100 cm and is soft and poorly compacted in areas without forest
cover. Caribou select old growth forests in the Larch/Western white
pine/Western fir-spruce zonal interface (25). The forest vegetation
zones occupied are more clearly defined by Krajina (23) as the
interior cedar-hemlock (ICH) biogeoclimatic zone and the low portion
of the Englemann spruce subalpine fir (ESSF) biogeoclimatic zone. The
elevations encompassed by these zones shows some variation over the
range of woodland caribou habitat in British Columbia. Simpson et al.
found caribou in an elevational range from 750 m to 1,900 m in
November and December with a steady downward trend from October to
December (15). The valley bottom elevation in that area is 570 m and
caribou moved almost to the valley bottom. In the southern Selkirks
Scott and Servheen found that caribou rarely descend below 1,360 m
(18). The valley bottoms in the south Selkirks are warmer, dryer and
have much greater human development. These caribou have the potential
to move below the snowline during early winter in the south Selkirks
while further north they do not, and yet they remained on snow at
elevations of 1,360 to 2,120 m. The reasons for these differences in
early winter habitat selection remain undetermined. Simpson et al.
Habitat Associations - 2 (15) consider early winter to be the most difficult season of the year
for caribou as food is most difficult to obtain and movement is most
difficult due to snow softness.
Caribou in some areas actively seek active logging for the forage
it provides on felled trees (15).
Disturbance of early winter habitats through logging or wildfire
has the potential to impact caribou severely. Extensive and complete
removal of old growth and mature forest would be detrimental to
caribou. The degree of removal which can be accommodated remains
undetermined. It is clear that forest management must insure the
continuous presence of old growth forests on some portion of the
caribou range. Scott and Servheen (18) recommend that a minimum of
25% of the old growth cedar-hemlock forest be maintained and that a
minimum of a 200 year forest rotation be established. They also
recommend the smallest feasible cut block sizes.
Late winter normally begins in January as the snow pack builds to
greater depths and it becomes more firm as the result of wind action,
thaws, and characteristic snow metamorphosis. A reasonably firm, deep
snow pack at high elevations permits caribou to forage on arboreal
lichens up to 6 m above ground level (17,23). Typical caribou habitat
during this period is the Western fir-spruce/Alpine tundra (25)
interface or Englemann spruce-subalpine fir (ESSF) parkland (15,26).
Caribou move to this habitat type at elevations of 1,700 m to 2,100 m
when the snow is firm enough, and remain there until mid-April or
later. Little variation has been found in habitat selection during
this period across the range of woodland caribou (15,17,18,23,27).
The main differences which can be observed are in the topographic
nature of the habitats. Extensive late winter habitats in the Wells
Gray Park area of B.C. are moderately sloping plateaus while the north
and south Selkirk mountain herds tend to occupy rounded ridges,
ciques, and gently sloping passes of much less area. Throughout the
Monashee, Selkirk and Purcell mountains of southern B.C. caribou are
typically found on the most gently sloping areas available at
elevations between 1,700 and 2,100 m during late winter (28). Simpson
et al. (15) considered late winter to be a period during which caribou
condition declined steadily but did not consider it to be as taxing a
period for caribou as early winter. Arboreal lichens are readily
available and make up almost 100 percent of the diet during this
period (14,15,17,18,29,30).
A need for arboreal lichens in late winter makes the forests
which the lichens grow upon most important. Lichen volume on a forest
is variable and seems dependent on slope position, aspect, slope, and
elevation (31). Winter lichen availability is further influenced by
snow depth and icing conditions on the trees. However, it is clear
that trees must be present as a substrate for the lichens, that they
must be large enough to reach well above the 2 to 3 meter snow pack,
and that they must be old enough to support a significant volume of
lichens. This means mature and old growth forests must be maintained.
Logging or wildfires can eliminate the needed forest. It is not yet
clear how much forest must be maintained but current lichen biomass in
the south Selkirk mountains is considered adequate to meet the needs
of the present caribou population (32).
Spring habitat selection patterns are variable over the range of
woodland caribou habitats in British Columbia but all seem to shift to
lower elevations. Scott and Servheen 918) found that the Selkirk
caribou selected elevations between 1,200 and 1,810 m and moved into
areas which had little snow, and were greening up. In the Revelstoke
area most caribou shifted right to the valley bottom at 500 m as soon
as the valleys greened up (15,17). In the north Thompson area,
caribou move down to mid-elevation areas to take advantage of early
green up but also do not move to valley bottoms (33). Simpson (34)
Habitat Associations - 3 found that habitat selection during spring favored open forested
habitats with good visibility, apparently to minimize predation by
grizzly bears. Scott and Servheen felt the Selkirk caribou population
selected spring habitat primarily for the availability of early
growing green forage (18,32). These were habitats on southwest
aspects, as well as valley bottoms and lower slopes. Forest stands
with 35% or less cover, or unforested areas were also preferred. It
is noteworthy that the Selkirk caribou did not move to low elevation
valley bottoms at 500 to 800 m where green up was much earlier,
although those valleys are present within normal travel distances for
caribou.
Habitat types occupied during spring include the Western white
pine, Western fir-spruce, larch, and lodgepole pine types or the lower
portion of the Englemann spruce subalpine fir zone (ESSF) and the
interior cedar hemlock (ICH) zone (17,18,24).
Scott and Servheen (18) recommend that spring habitats be managed
to maintain 25% of the zone in preferred habitat using short stand
rotation lengths, leaving residual trees, and opening up all but
pockets of old growth cedar-hemlock. Simpson considered heavy slash
loads following logging to be detrimental to spring caribou habitat
and recommended clearing after logging (34).
Calving habitat is distinct from spring or summer caribou
habitats. Female caribou shift away from other caribou prior to
calving and travel to habitats at high elevations, near ridge tops, on
steep slopes (17,18,32). These are normally western fir-spruce types,
and snow is normally still present. Forests in these areas were
mainly undisturbed, moderately-canopied (16-75%) stands of medium to
large diameter trees. Englemann spruce, subalpine fir and whitebark
pine were the main species present (32). The habitat selection
observed during calving is considered to be a predator avoidance
strategy (17,34). Grizzly bears are the main predators to be avoided
during this period in the south Selkirk Mountains, while black bears
may also be important. Wolves are important predators on caribou in
the Quesnel highlands of British Columbia (27) and may have been a
historically important predator in the Selkirk range.
It is not clear whether calving habitats are likely to be
adversely impacted by human activities. They are generally poor
forest sites and are unlikely to be logged. Wildfire has the
potential to kill the trees over extensive calving areas.
Summer, fall, and rut habitats are the least clearly selected of
any season as abundant forage is available. Caribou seem to select
areas which provide an abundance of succulant forage for as long as
possible. These sites were on moist north and east aspects (18,32).
Western fir-spruce habitats are primarily used, although a range of
other habitats are used during incidental movements between areas.
In general, aquatic habitats are not specifically selected with the
exception of palustrine emergent systems (bogs, fens, sedge meadows)
which are used for foraging. Agricultural land is not commonly used
or selected by caribou but caribou have occasionally been observed on
hay fields or pastures. The significance of this use is not known.
It is possible that the use of hayfields and pasture reflects a
historic use of open natural meadows which have been replaced by
agriculture. Riverine systems are also used but generally only in
passing from one area to another.
Logged and wildfire areas are commonly used during summer as they
provide abundant forage. No specific habitat disturbance is likely to
have a major impact on caribou survival during summer.
The range of habitats used by woodland caribou across British
Columbia and into Idaho and Washington results in variation in habitat
selection (35). Caribou demonstrate adaptability, with changes in
diet seasonally and across their range, however, they are well adapted
Habitat Associations - 4 to a specific range of conditions. High elevations appear to be
preferred throughout the year as caribou are seldom found below 600 m
in B.C. (35). Forested habitats are preferred throughout the year,
although the amount and type of forest varies from season to season.
Mature to old growth forests are needed as they supply the growth
substrate for lichens needed during winter. The best woodland caribou
habitats in B.C. are generally rolling plateaus or gently sloping
mountain tops which support extensive subalpine forest. More steeply
sloping mountains and ridges such as found in the Selkirk and Purcell
mountain ranges support smaller and more scattered caribou populations
(35).
Habitat Associations - 5 (DRAFT) - Food Habits
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
FOOD HABITS
TROPHIC LEVEL -
HERBIVORE
LIFESTAGE FOOD FOOD PART
General Vascular Plants-Submerged Nonwoody
General Poaceae
General
General Evergreen Shrubs-Leaves/Twigs
General Deciduous Trees-Leaves/Twigs
General Coniferae
General Vascular Plants-Emergent Nonwoody
General Myxomycophyta
General Forb Leaves/Stems
General Forb Flowers/Fruit/Seed
General Deciduous Shrubs-Leaves/Twigs
General Deciduous Shrubs-Flowers/Fruit/Seed
General Vascular Plants-Submerged Nonwoody
General Poaceae
General
General Evergreen Shrubs-Leaves/Twigs
General Deciduous Trees-Leaves/Twigs
General Coniferae
General Vascular Plants-Emergent Nonwoody
General Myxomycophyta
General Forb Leaves/Stems
General Forb Flowers/Fruit/Seed
General Deciduous Shrubs-Leaves/Twigs
General Deciduous Shrubs-Flowers/Fruit/Seed
Food Habits - 1 (DRAFT) - Environment Associations
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G Inland Wetlands: Bogs
G Terrestrial Features: Downed logs
G Inland Wetlands: Bogs
G Terrestrial Features: Downed logs
Environment Associations - 1 (DRAFT) - Life History
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
LIFE HISTORY
FOOD HABITS:
The unique feature of woodland caribou, (Rangifer tarandus
caribou) food habits in the mountainous deep snow region of British
Columbia, Idaho, and Washington is their reliance on arboreal lichen
species for winter food (14,15,17,18,23,32,33). The lichen species
Alectoria sarmentosa and Alectoria jubata are almost their exclusive
diet from October until May.
Food habits of woodland caribou are seasonal in nature as a
result of the deep snow. Arboreal lichen makes up most of the diet
during winter once the snow covers other species. Fecal food habit
studies indicate that a moderate proportion of the diet consists of
spruce and fir needles during winter, but this is believed to be the
result of incidental ingestion while feeding on lichens (17,18).
As snow leaves the ground, species which green up early are
selected. Luzula hitchcockii growing in snow free tree wells is one
of the earliet species to begin growth and is well used (18,32).
During spring and summer caribou used huckleberry (Vaccinium
membranceum) leaves and mountain Valeriana (Valeriana stichensis) much
more than any other forage (18).
A few species were selected during early growth stages in spring
(Alnus sinuata buds, Epiolobium angustifolium, Lonicera utahensis,
Gymnocarpium dryopteris, and Erythronium grandiflorum). Other species
were preferred but were rare include Disporum hookeri, Smilacena
racemosa, Streptopus amplixifolius, and Sorbus scopulina (32).
Simpson (34) found sedges (Carex), horsetails (Esquisetum), Streptopus
amplexifolius, Vaccinium membranaceum, and grasses (Graminae) were the
primary spring forages, reflecting a greater use of riparian meadows
in the Revelstoke area, particularly prior to flooding of the valley
bottom by the Revelstoke Dam.
Other researchers found that caribou fed primarily on herbaceous
vegetation, mushrooms, shrub leaves, grasses, sedges, and soft shrubs
(19,23,36).
In fall, as snow began to accumulate and succulent green forage
became rare caribou would "crater" in 20-30 cm of snow for evergreen
forbs and shrubs such as Pachistima myrsinites, Pyrola spp., Tiarella
tifoliata, Viola orbiculata, Rubus pedatus and Lycopocium spp. (18).
Extensive cratering was not observed in the south Selkirks (18) nor
has it been observed in the Revelstoke area (17). However, woodland
caribou in west central Alberta crater extensively for terrestrial
lichens (37). Snow accumulations in those areas are much less, seldom
exceeding 1 m.
The shrublike or hairlike growth form of arboreal lichens results
in a relatively uniform forage which is entirely available to caribou.
Where forage is in short supply all available lichen is consumed along
with some incidental feed on the substrate leaves or branches. In
summer the caribou forage primarily on the leaves and buds of plant
species. Little browsing of woody stems occurs (06).
HOME RANGE/TERRITORY:
Woodland caribou do not establish or defend a territory and in
some areas do not occupy what can truly be considered a home range.
Life History - 1 (DRAFT) - Life History
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
In the Selkirk mountains caribou occupied a core area and shifted to
seasonal habitats at a variety of locations, but no overall movement
out of one core area was observed over 3 years (18,32). Bull
movements were more far ranging than cow/calf group movements. In
other areas, however, seasonal habitats were occupied, but no defined
core area was established. In the northern Selkirks a more wandering
pattern of seasonal habitat selection was observed (17,27).
Seasonal home ranges in the southern Selkirk mountains were
highly variable between individuals but in general are smallest during
early winter (25.7 sq. km) and spring (27.0 sq. km) and largest during
late winter (44.2 sq. km). Annual ranges for 2 cows averaged 111.2
sq. km (18). In the Revelstoke area annual ranges varied from 151 sq.
km to 978 sq. km over a 4 year period (17).
Bull home ranges in the south Selkirk Mountains were larger than
cow home ranges both seasonally and annually. The greatest difference
appeared to occur in early winter when bull seasonal ranges were from
35.4 to 79.5 sq. km. Annual ranges for 2 bulls were 245.1 and 253.7
sq. km (18).
Social groups of woodland caribou are loose and variable. Stable
groups of 1 to 20 animals will form during some seasons, particularly
late winter, but often break up to establish other groups (17,18,32).
Cows, calves, and immature bulls occur in mixed groups. Mature bulls
are sometimes found with cow/calf groups, sometimes with small bull
groups and sometimes alone (05,17,18,32).
PERIODICITY:
Daily periodicity of caribou has not been examined closely.
During most seasons feeding and travel movements take place day or
night. During early winter on areas undergoing logging a crepuscular
shift into the logged areas to feed occurs but it may be closely
related to the end of intense daytime logging activities. Further
research is required.
MIGRATION PATTERNS:
Woodland caribou do not demonstrate the marked annual migration
pattern exhibited by barren-ground caribou but annual habitat
selection shifts or elevational migrations do occur. Early winter
habitats tend to be at lower elevations, and as late winter snow
compaction occurs the caribou shift upward onto parkland habitats. As
spring green-up begins at lower elevations a downward shift occurs to
seek green forage. A gradual upward shift occurs from spring to
summer as the snowline rises. As winter approaches the caribou again
shift down onto early winter habitats. A distinct upward movement
takes place before calving in June as cows leave the spring green-up
areas and move to calving habitats on high, snowy ridges. They
usually return to green-up areas 2 to 3 weeks after calving (05,15,17,
18,21,23,34,37).
In some areas of cental British Columbia and Alberta more
migratory behaviour occurs as caribou shift between summer and winter
ranges. These caribou occupy winter ranges with significantly less
snow accumulation than is normal in caribou habitats further south
(35,36).
Life History - 2 (DRAFT) - Life History
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
COVER/SHELTER REQUIREMENTS:
Cover requirements of woodland caribou are seasonally dependent,
but may be more related to cover as a food source or substrate than
for shelter. In early winter mature or old growth Englemann spruce
(Picea englemannii), subalpine fir (Abies lasiocarpa), red cedar
(Thuja plicata), and hemlock (Tsuga heterophylla) stands intercept
snow and result in greater snow compaction beneath the trees. This
cover is selected for its snow compaction and easier travel (15,18),
in addition to the lichen food supply resulting from lichen litter
fall and windthrown trees.
Scott and Servheen found the Selkirk caribou population seemed to
select forest stands of 53 cm dbh or greater during early winter (18).
Preferred overstory canopy cover in these forests ranged from 46 to
75%. In late winter open forest canopies are preferred (15,18).
Selkirk caribou preferred forested habitats with timber sizes classes
from 20 cm to 53+ cm, and overstory cover of 46 to 100%. Caribou
often bedded down in the small openings during this period and showed
no evidence of seeking shelter from wind or cold (15).
Spring, summer and fall cover selection was more variable than
during winter in the south Selkirks. Simpson (24) has identified
spring habitats as riparian meadows, cedar-hemlock forest, and
Englemann spruce-subalpine fir forest, with low visual and movement
obstruction values as to a predator avoidance strategy. Grizzly bears
are commonly found on the most productive sites but sight distances
are poor there and caribou are believed to be selecting sites with
less forage but better grizzly avoidance characteristics. Open
forests or logged areas are used throughout the summer and fall. No
shelter or cover requirements have been identified during this period.
REPRODUCTIVE SITE REQUIREMENTS:
As autumn approaches a general habitat selection pattern for
moist forest sites becomes more pronouned as caribou selected gently
sloping secondary stream bottoms at lower elevations. The rut takes
place while caribou occupy these habitats in the 1,360 to 1,800 m
elevation range. Logged and unlogged sites are used but the
preference appears to be for unlogged old growth forest sites (18).
These habitats are not selected for the rut; they appear to be
selected as suitable forage sites during that period.
Calving habitat is very distinct, as just prior to calving
females shift away from spring habitats to isolated high elevation
forests at or near ridge tops (17,18). In the south Selkirks caribou
selected southwest aspects on upper slopes or ridge tops above 1,810 m
elevation (18). A wide range of pole-sized to old growth forest types
were used (18). This habitat selection pattern is believed to be a
predator avoidance strategy (17,18). Cows remained alone during this
period. Cow-calf pairs remained in these habitats until mid-July
unless the calf died, in which case the cow immediately rejoined the
main caribou groups (18). No particular habitat selection or
behavioural patterns have been observed once calving habitats have
been occupied, however data on this subject is scarce as research
during this period is difficult.
REPRODUCTIVE CHARACTERISTICS:
Life History - 3 (DRAFT) - Life History
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
Little research and secretive habits have resulted in the present
scarcity of information on woodland caribou reproductive
characteristics. Some characteristics may be similar to barren ground
caribou (Rangifer tarandus groenlandicus) and the literature on that
subspecies has been drawn upon.
Scott (18) documented a movement by mature bull caribou to the
area utilized by cow and calf groups in mid-Septimber to mid-October.
Researchers have not documented rut behaviour. Bulls do not defend
a harem and they are believed to breed females as they become
receptive. No pair bonding occurs. Bulls are believed to assert
dominance through displays and fights, and dominant bulls are believed
to do most of the breeding as do the barren ground caribou bulls (39).
Age of sexual maturity among woodland caribou is also unknown. It is
likely to range between 18 and 42 months, for females, as it does for
barren ground caribou. Males likely mature at approximately 18 months
(39). Maximum breeding age is unknown. The receptive period of the
female is not specifically known but is likely to occur once per year
in late October to early November. It is likely that second and third
receptive periods follow the first by 10 to 12 day periods as it does
in northern caribou (40). The gestation period of woodland caribou in
this area has not been investigated but is likely to be very close to
the 227 to 229 day period found in other Rangifer species (36). Only
one calf is normally born per year and twinning is very rare (36).
Calving has not been observed as cows become very secretive during
calving and move to inaccessible habitats. Calves are believed to be
precocial as they are for other Rangifer species (39). If the barren
ground example is correct, calves will be able to walk and follow the
mother 1 to 3 hours after birth and will be able to travel well at 1
day (07).
PARENTAL CARE:
Little is known about parental care among woodland caribou.
Calves are born in early June (15,18) and are believed to be precocial
as are barren ground caribou (41). Barren ground caribou calves begin
grazing within 2 to 4 days after birth and graze almost exclusively
after only 3 weeks (39). Whether this is the case among woodland
caribou is unknown. Calves normally remain with the same group
through the winter, although little attachment to the mother is
believed to exist.
Woodland caribou cow/calf pairs normally remain away from other
caribou for a period of one to three weeks at the time of birth. This
is believed to occur to provide protection from predation during the
early period of the calf's life as the cow selects habitats which are
relatively safe from predators (15,18). Behavioural protection of
woodland caribou offspring is unresearched.
Bull caribou may remain with cow/calf groups or may wander
solitarily or with other bulls. They appear to play no role in
maintenance or protection of offspring.
POPULATION BIOLOGY:
Across the range of woodland caribou in British Columbia,
Washington, and Idaho several limiting factors act on the populations
but researchers are not in agreement on the importance which should be
Life History - 4 (DRAFT) - Life History
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
placed on each factor. Long-term food supplies are a specific concern
in some areas of B.C. as logging threatens to eliminate old growth
forests on caribou wintering areas (35). In some areas wolves, and
possibly grizzly bears may be influencing populations (35). Hunting
and poaching are considered to have caused current low populations in
much of southern B.C. Natural catastrophies such as avalanches are a
significant mortality factor in some areas (17). Human development
may have reduced caribou use of main valleys. Highway mortality in
the south Selkirk population may help limit its growth (41). The
meningeal worm (Paralophostrongylus tenuis) is not believed to be
present in B.C. and therefore, is not a factor as it is in eastern
Canada (08). Scott and Sevheen (18) considers illegal hunting and
poor neonatal calf survival to be the cause of current low caribou
numbers in the Selkirk population. In the Revelstoke area the
population is considered to be increasing. Hunting and poaching in
that area has been controlled, and predators are not considered to be
a factor as caribou successfully avoid the grizzly bears, the only
significant predator at this time (17).
Detailed information on survival rates, mortality rates, rates of
increase and other life table calculations are not yet available.
Scott and Servheen (18) found low calf survival rates as only 3 to 5
calves were produced from a potential 7 to 9 annually, resulting in
calf survival rates of 25 to 50% annually. At least 45% of adult cows
failed to produce calves, primarily due to a high neonatal death rate.
Simpson et al. (17) found that calves made up 18.9% to 25.8% of the
total population over the course of the study as compared to 11% to
18% in south Selkirks (18). Seip and Hebert (27) found 100% pregnancy
rates in the Quesnel Highlands caribou. Neonatal calves per 100 cows
ranged from 38.5% to 55% while spring calf recruitment rates had
dropped to 3 to 11 calves per 100 cows. These calf mortalities were
partly attributed to predation.
Adult mortality rates have been high as adults seem to be
vulnerable to some predation by grizzly bears and wolves, to
accidental death due to environmental hazards such as avalanches, and
to illegal hunting (17,18,27). Bergerud (36) indicated that natural
adult caribou mortality rates ranged from 5 to 13%. In a very small
sample, Simpson et al. (17) observed a mortality rate of 25% - 33% as
a result of natural causes. Lack of data has prevented calculation of
population rates of increase to date. Scott and Servheen (18)
regarded the Selkirk population as stable or declining, Simpson et al.
(17) regarded the Revelstoke population as stable or increasing. Sex
ratos among adults in the Revelstoke population appear to be about
equal, while subadults remained unidentified due to classification
difficulty (17). In the south Selkirks the population composition
ranged from 11 to 18% calves, 41 to 50% cows, 21 to 37% immature bulls
and 11 to 15% adult bulls (18).
Recovery potential in the south Selkirks is not fully understood.
Bergerud (36) regarded caribou populations with > 15 percent
calves in spring as increasing. There appears to be a fluctuation of
calf survival around 15% but natural and human induced adult mortality
is high. Turnover rates and longevity in these caribou populations
remains researched.
Population density objectives for the south Selkirk populations
Life History - 5 (DRAFT) - Life History
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
have not been set at this time. A recovery goal of 100 animals was
set (05) and current population densities in the occupied habitat are
1/7.5 sq. km (18). In the Revelstoke area a population density of
1/23 sq. km was observed (17) while in the Quesnel Highlands densities
are about 1/10 sq. km (27).
SPECIES INTERRELATIONSHIPS:
Woodland caribou calves are vulnerable to predation for the first
days of life. The primary predators likely to prey on caribou are
wolves (Canis lupus), grizzly bears (Ursus arctos horribilus), and
black bears (Ursus americanus). A predator avoidance strategy
employed by caribou during calving is believed to greatly reduce
vulnerability to predators (17,18) but lack of data on immediate post
parturition causes of calf mortality leaves the question unresolved.
Predation on adult caribou seems to be a rare occurrance as
little evidence of predation has been collected during woodland
caribou studies. In the Revelstoke area a predator avoidance strategy
appears to be employed by caribou on spring ranges (17). Wolves are
believed to play a significant role in the life of woodland caribou in
the Quesnel Highlands (27). However, wolves do not presently inhabit
the south Selkirk Mountains and do not play any role in the survival
of caribou along the U.S. border. Moose (Alces alces), elk (Cervus
elaphus), white-tailed deer (Odocoileus virginianus), and mule deer
(Odocoileus hemionus) all inhabit woodland caribou habitat during
summer. No significant inter-relationships have been observed.
OTHER LIFE HISTORY DESCRIPTORS:
None known.
Life History - 6 (DRAFT) - Management Practices
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Adverse
Adverse Commercial Exploitation
Adverse Fire
Adverse Food Supply Reduction
Adverse Forest Alteration
Adverse Harvesting
Adverse Harassment/Vandalism/Indiscriminate Killing
Adverse Highway/Railroads
Adverse Inherent Reproductive Characteristics
Adverse Poaching
Adverse Predation
Adverse Rural Residential/Industrial Areas
Adverse Sport Hunting/Fishing
Adverse Vegetation Composition Changes
Beneficial Stocking captive-reared wild-strain animals
Beneficial Suppressing wildfire
Beneficial Restricting/regulating human disturbance of populations
Beneficial Maintaining Later Stages of Succession
Beneficial Restricting Poaching
Beneficial Selective Thinning
Beneficial Transplanting wild animals
Existing
Existing Commercial Exploitation
Existing Fire
Existing Food Supply Reduction
Existing Forest Alteration
Existing Harvesting
Existing Harassment/Vandalism/Indiscriminate Killing
Existing Highway/Railroads
Existing Inherent Reproductive Characteristics
Existing Poaching
Existing Predation
Existing Rural Residential/Industrial Areas
Existing Sport Hunting/Fishing
Existing Vegetation Composition Changes
COMMENTS ON MANAGEMENT PRACTICES -
No specific reason for woodland caribou population declines has
been identified and agreed upon by caribou researchers (08).
Availability of adequate winter forage, caribou habitat changes
resulting from logging and fire, market hunting, sport hunting,
poaching, construction of roads in caribou habitat, agricultural
development and grazing, rural residential development, and predation
all likely have influenced populations with a combination of factors
likely resulting in population elimination or reduction.
In the Selkirk Mountains Layser (10), Miller (11), and Stevenson
(12) identified lichen availability in winter as a key factor in
caribou survival. Availability of lichens is believed to have been
Management Practices - 1 (DRAFT) - Management Practices
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
influenced by logging and wildfire. Disturbance of early winter
habitats through logging or wildfire has the potential to impact
caribou severely. Extensive and complete removal of old growth and
mature forest would be detrimental to caribou. The degree of removal
which can be accommodated remains undetermined. It is clear that
forest management must insure the continuous presence of old growth
forests on some portion of the caribou range. Scott and Servheen (18)
recommend that a minimum of 25% of the old growth cedar-hemlock forest
be maintained and that a minimum of a 200 year forest rotation be
established. They also recommend the smallest feasible cut block
sizes. Hunting, poaching and collisions with motor vehicles have been
a major cause of caribou mortality during the last 3 decades (07,13,
14,15). Bergerud (16) attributed population declines in southern B.C.
to increased hunting and poaching which resulted from increased
access.
Predation has been considered a major source of mortality.
Wolves (Canis lupus) are absent from caribou ranges in most of
southern British Columbia and are not currently a factor in predation.
Black bear (Ursus americanus) and grizzly bear (Ursus arctos
horribilus) are common in the area but are not considered to be a
significant mortality factor (17, 18). Scott (18) concluded that the
Selkirk caribou herd population declines may be the result of
unidentified natural calf mortality, and illegal hunting.
Caribou have been extirpated from most of their historical range
in the United States. Legal and illegal market hunting combined with
natural mortality rates are likely to have reduced populations (19).
Survival of small populations was likely jeopardized by habitat losses
due to wildfire, and logging as well as isolation due to extensive
agricultural development, rural residential development and
transportation corridors in major valleys separating caribou habitats.
Meningeal worm (Parelaphostrangylus tenuis) may be a primary
cause of caribou mortality in eastern Canada and the eastern United
States where white-tailed deer (Odocoileus virginianus) and caribou
ranges overlap. However, menengeal worm has not been identified in
white-tailed deer or caribou in western North America (09).
A future threat to woodland caribou may be gene pool narrowing as
a result of geographical isolation (05). Research to determine the
current genetic diversity in the existing Selkirk caribou herd and
other caribou populations began in 1985 (20). All of the existing
threats to caribou will continue in the future. Efforts are being
made to reduce human induced mortality factors and habitat or forage
limitations.
APPROVED PLAN:
U.S. Fish and Wildlife Service. 1985. Selkirk Mountain Caribou
Management Plan. U.S. Fish and Wildlife Service, Portland, OR.
118 pp.
The primary objective in the Selkirk Mountain Caribou Management
Plan/Recovery Plan is to increase the number of Selkirk Mountain
caribou from a declining herd of about 28 to a viable population
through identifying actions and implementing plan strategies.
Management Practices - 2 (DRAFT) - Management Practices
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
Criteria for declassifying to threatened status and delisting the
population have not been developed. An intermediate goal of 100
animals was set by several of the involved agencies. Current
population densities in the occupied habitat are 1/7.5 sq. km (18).
The recovery activities necessary to accomplish the objective
are:
1. Establish caribou population goals for maintenance of a viable
population by land management unit.
2. Determine management objectives for caribou habitat to provide
for the goal population.
3. Identify the information needs for research and develop strategy
for securing the information. Selective thinning and uneven-aged
timber management systems may be valuable tools in maintaining or
improving caribou habitat.
4. Evaluate and implement methods to increase population by the
transplanting surplus animals from other areas to the existing herd,
establishing a second herd (possibly through captive breeding for
reintroduction), protecting habitat, prevent fires, close roads to
public access, and reduce hunting in habitat, and preventing
accidental or malicious mortalities.
5. Implement public information and education program to gain public
support and to reduce poaching, highway mortalities, and other
decimating factors.
6. Implement management plan.
Management Practices - 3 (DRAFT) - References
Species CARIBOU, WOODLAND
Species Id ESIS051007
Date 14 MAR 96
References
***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE *****
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Columbia. Provincial Mus. Handbook No. 11, Victoria, B.C. 414 pp.
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America, New York. 780 pp.
03 Allen, J.A. 1903. Mammals collected in Alaska and Norther British
Columbia by the Andrew F. Stone expedition of 1902. Bull. Amer.
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04 Banfield, A.F. 1961. A revision of the reindeer and caribou,
genus Rangifer. Natl. Mus. Can. Bull. 177. 137 pp.
05 U.S. Fish and Wildlife Service. 1985. Selkirk Mountain Caribou
Management Plan. U.S. Fish & Wildl. Serv., Portland, OR. 118 pp.
06 Neuner, A.M. and T.J. Berger. [n.d.] Checklist of mammals of the
United States and the U.S. Territories. U.S. Fish & Wildl. Serv.
Coop. Agreement #14-16-0009-1038. 31 pp.
07 Freddy, D.J. and A.W. Erickson. 1972. Status of the Selkirk
mountain caribou. IN: Proceedings, First International
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08 Bergerud, A.T. 1974. Decline of caribou in North America
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09 Burke, T.E. and D.R. Johnson. 1984. A report on the potential for
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10 Layser, E.F. 1974. A review of woodland caribou of northeastern
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11 Miller, D. 1975. Unpubl. rep. Progress Report, Selkirk mountain
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12 Stevenson, S.K. 1979. Effects of selective logging on arboreal
lichens used by Selkirk caribou. Fish and Wildlife Report R-2,
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13 Bloomfield, M.I. 1980. The impact of development, settlement, and
associated activities on mountain caribou in Central British
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herd 1973. M.S. Thesis, Univ. Idaho, Moscow. 132 pp.
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of caribou (Rangifer tarandus caribou) in the mountains of southern
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17 Simpson, K., K.B. Hebert, and G.P. Woods. 1984. Unpubl. rep.
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Columbia Mountains of British Columbia. B.C. Fish & Wildl. Rep.,
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18 Scott, M.D. and G. Servheen. 1985. Caribou ecology. Job
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19 Evans, H.F. 1960. A priliminary investigation of caribou in
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20 Chesser, R.K. 1984. Genetics of the Selkirk caribou herd.
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24 Simpson, K. 1983. Arboreal lichen growth rates on subalpine fir
(Abies lasiocarp) north of Revelstoke, B.C. B.C. Fish and Wildlife
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25 Garrison, G.A., A.J. Bjugstad, D.A. Duncan, M.E. Lewis, and D.R.
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28 Woods, G.P. Unpubl. data. Fish and Wildlife Branch, 310 Ward St.,
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29 Munro, J.A. 1945. Preliminary report on the birds and mammals of
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30 Hamer, D. 1974. Distribution, abundance and management
implications of grizzly bear and mountain caribou in the Mountain
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31 Detirck, R.W.T. 1984. Unpubl. rep. Arboreal lichen biomass
available to caribou, Selkirk Mountains, northern Idaho. Idaho
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32 Scott, M. and G. Servheen. 1984. Caribou ecology. Idaho Dept.
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33 Antifeau, T. 1980. North Thompson River mountain caribou study.
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34 Simpson, K. 1986. Impacts of a hydro-electric reservoir on
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Species Id ESIS051007
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caribou (Rangifer tarandus caribou) in west central Alberta,
1979-1983. Alberta Energy & Natl. Resour., Fish & Wildl. Div.
203 pp.
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and Northern Develop. Queen's Printer, Ottawa, Ont. 340 pp.
40 Dauphine, T.C., Jr. 1976. Biology of the Kaminuriak population of
barren-ground caribou. Part 4. Growth, reproduction, and energy
reserves. Canadian Wildlife Service Report 38. 71 pp.
41 Johnson, D.R. 1976. Mountain caribou, threats to survival in the
Kootenay Pass region, Brittish Columbia. Northwest Sci.
50(2):97-101.
42 Meatte, D.S. 1986. Evidence for the historic occurrence and
aboriginal utilization of caribou in southern Idaho. Paper
presented at the 13th Ann. Conf. Idaho Archaeological Soc., College
of Southern Idaho, Twin Falls, Sept. 27, 1986. Available from:
Danil S. Meatte, Dept. of Anthropology, Sociology, and Social Work,
Idaho State Univ., Pocatello. 8 pp + 1 map.
43 Hall, E.R. 1981. The Mammals of North America. John Wiley &
Sons, Inc., New York. 1175 pp.
***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY *****
01 Servheen, Greg. 1985. Personal communication. Idaho Game Dept.,
Bonner's Ferry, ID 83805.
02 Scott, M.D. and G. Servheen. 1985. Caribou ecology. Job
Completion Report, Idaho Dept. of Fish and Game. 136 pp.
03 Evans, H.F. 1960. A priliminary investigation of caribou in
northwestern United States. M.S. Thesis, MT State Univ., Bozeman.
04 Mette, D.S. 1986. Evidence for the historic occurrence and
aboriginal utilization of caribou in southern Idaho. Paper
presented at the 13th Ann. Conf. Idaho Archaeological Soc., College
of Southern Idaho, Twin Falls, Sept. 27, 1986. Available from:
Daniel S. Meatte, Dept. of Anthropology, Sociology, and Social
Work, Idaho State Univ., Pocatello. 8 pp + 1 map.
05 Anderson, E. and J.A. White. 1975. Caribou (Mammalia, Cervidae)
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