Abstract
Bounty records from Montana (1902–1930) were used to better understand spatial and temporal response of wolves (
Canis lupus) and cougars (
Puma concolor) to an eradication program. Number of payments for wolves declined from a high of 4116 in 1903 to 0 by 1928. Numbers of cougar pelts fluctuated, but generally declined from 177 in 1908 to 2 in 1930. An inverse spatial relationship
existed between density of payments for wolves and cougars. Payments for wolves were greatest in the Prairie ecoregion, while
payments for cougars were greatest in the Montane(MOUNTAINS). The ratio of pup to adult wolves also was greatest in the Prairie ecoregion and
least in the montane. If distribution and frequency of bounty payments were a reflection of the distribution and abundance of
wolves, populations were highest and most productive in areas where they currently do not occur.--note that where cougars and wolves occupy the same region, cougars tend to occupy higher Country where their stealth and ambush style hunting is optimized...........wolves as a coursing(chasing) predator prefer the open flatlands..............this is how the two predators minimize their conflicts with each other--blogger Rick
Once two of the most widely distributed large carnivores
in the world, wolves (
Canis lupus) and cougars (
Puma concolor), were nearly eradicated in the western US as a result of habitat loss and predator control
programs during the 20th century (Mech, 1970; Young
and Goldman, 1944). Central to the success of predator
control programs were economic incentives in the form
of payments (bounties) to hunters or trappers in exchange
for animal parts or pelts. Records of bounty
payments provide a novel source of data regarding distribution
and relative abundance of wolves and cougars,
and response of large carnivores to human-caused exploitation.
Improved understanding of where ancestral
populations existed and their response to exploitation
provide a historical perspective to management of large
carnivores such as wolves and cougars, and may aid
current conservation efforts for these species (Berger and
Wehausen, 1991; Fuller and Kittredge, 1996).
The first territorial bounty legislation aimed at wolves
and cougars in Montana was enacted in 1883 (Curnow,
1969). Bounty payments occurred until 1887, when the
bounty law was repealed. A new bounty program was
reinstated in 1891. By 1930, wolves were eradicated from
Montana, and few cougars remained (Bangs et al., 1998;
Young and Goldman, 1944, 1946).
County Treasurers kept careful
records of the eradication program through meticulous
ledgers of bounty payments. These ledgers provide a
source of harvest data, which may provide insight into
historical exploitation and eradication of two species of
large carnivores.
The goal of this study was to increase understanding
of spatial and temporal dynamics of exploitation of
large carnivores, and identify implications of this understanding
for current management of these species.
Montana has a land area of 381,086 km2, and averages approximately 870 km from east to west and 450
km from north to south (Fig. 1). Elevations range from
546 m where the Kootenai River exits the state in the
northwest corner to 3898 m in the Beartooth Mountains
of south central Montana. The western third of Montana
is mountainous and the eastern two-thirds are a
mix of undulating cleft prairie interspersed with rivers
and isolated mountain ranges of the Missouri and Yellowstone
watersheds.
All counties of Montana were assigned to one of
three main ecoregions, the Montane, Intermountain,
and Prairie ecoregions (Riley, 1998). The Montane ecoregion
was comprised of Northern Rocky Mountain
Steppe–Coniferous Forest–Alpine Meadow Provinces
(McNab and Avers, 1994) within the Columbia Basin of
northwestern Montana. Southwestern Montana and the
Rocky Mountain East Front were classified into an Intermountain
ecoregion, which contained a mix of Middle
Rocky Mountain Steppe–Coniferous Forest–Alpine
Meadow and Southern Rocky Mountain Steppe–Open
Woodland–Coniferous Forest–Alpine Meadow Provinces.
The remainder of the state is predominately in the
Prairie ecoregion, comprised of Great Plains–Palouse
Dry Steppe Province.
Bounty harvest records from 1902 to 1930, originally
recorded by county clerks, were transcribed from ledgers
archived in the Montana State Library in Helena. Record
books from 1924 were missing. Records represented
number of bounty payments for cougar, adult
wolf, and wolf pup pelts submitted by individuals in the
county for which the animals were killed. Cougar
bounty prices changed from $7 in 1902 to $10 in 1905.
Adult wolf bounty prices changed from $5 in 1902 to
$10 in 1906 and $15 dollars in 1912. Bounty price for
wolf pups was $3 until 1904 and $4 thereafter.
We summarized the proportion of total wolf and cougar bounty payments made in each ecoregion.
Bounty records may provide a glimpse of historic
wolf and cougar ecology as well as population response
to human exploitation. However, caution need be exercised
when making inferences from the data. Some
patterns we observed may be an artifact of wolf and
cougar vulnerability to human harvest, or errors in reporting
bounty kills. Data are lacking on bounty hunter
effort, which might enable stronger inference between
bounty kills and actual populations. Another source of
potential bias is our inability to document the entire
history of wolf and cougar exploitation in Montana.
Bounty kills in Montana began in 1883, but only records
from 1902 to 1930 were retrievable. Despite these potential
sources of bias, we believe patterns from these
data may be valuable for conservation planning.
Number of payments declined rapidly from a peak
in 1903 until 1909 (Fig. 2). A temporary increase
in payments occurred from 1911 until 1914, after which
a decline occurred until wolves were extirpated. No
payments for wolf pelts were made in Montana after
1927.
Bounty payments for wolves were widespread across
Montana from 1902 to 1910, but were greatest in the
eastern half of the state. Through time, the
pattern of payments became more scattered across
Montana as payments were no longer reported in some
counties. Wolf kills from 1911 to 1920 were concentrated
in the middle of the state with brief, but relatively
high densities of kills localized in the northwest and
southeast. Small numbers of wolf pelts were presented in
a scattering of counties across most of the state during
the last 9 years wolves were present in Montana. One
exception was the Montane ecoregion, where no payments
were made after 1921. The last bounty payment
for wolves was recorded in Powder River County in
southeastern Montana.
The ratio of wolf pups to adults was greatest in the
Prairie ecoregion (Fig. 3). The ratio of pups to adult
wolves presented for payment never exceeded one in the
Montane (max
¼ 0:07) or Intermountain (max ¼ 0:57) ecoregions.
Cougar payments fluctuated from 1902 to 1922 with
peaks occurring at an interval of 3–4 years
Payments were greatest in 1908, when 177 pelts were
submitted for payment. By 1930, only two payments
were made for cougar pelts in Montana.
From 1902 to 1910, nearly all counties in Montana
had cougar pelts presented for bounty payment (Fig. 2).
The highest numbers consistently occurred in the
Montane and Intermountain ecoregions. By 1911, few
pelts were presented for payment in the eastern half of
the state. From 1925 to 1930, less than five pelts per year
were presented across the state. Numbers of cougar pelts
were more concentrated in the Montane ecoregion except
in the last 5 years when they were scattered across
the western half of the state.
Bison bison), s first bounty program both occurred Odocoileus virginianus) populations (Riley andMalecki, 2001). Historically, many of these valleys were inhabited
by wolf populations that may have discouraged cougars
from descending to lower elevations (Mussehl and Howell,
1971). If wolf populations continue to expand into
areas currently occupied by cougars, wolves may once
again restrict cougars to more mountainous and rugged
terrain. Research is needed to better understand dynamics
of potential competitive interactions between
wolves and cougars, and their implications for conservation.
Dynamics of early wolf and cougar eradication efforts in
Montana: implications for conservation
The spatial differences in abundance and productivity
we observed between wolves and cougars have several
implications for conservation. Wolves in the western US
currently are limited to mostly mountainous habitats of
Idaho, Montana, and Wyoming, on or adjacent to national
forests and national parks (Bangs and Fritts,
1996; Bangs et al., 1998). If historical distribution and
abundance of bounty payments were a reflection of
earlier distribution and abundance of wolves in Montana,
ancestral wolf populations likely were highest and
most productive in areas where they currently do not
occur. Localized pockets of high bounty yields were
observed in the Montane and Intermountain ecoregions.
However, these ecoregions did not sustain high yields
for as long as the eastern Prairie ecoregion. High apparent
productivity of wolves in the prairie was expressed
in the ratio of pups per adults presented for
payment, and this occurred despite high human-caused
mortality. Other ecoregions of Montana experienced
lower harvests, yet these ecoregions did not appear to
support sustained wolf productivity.
in 1883. Wolf populations probably switched
prey from bison to cattle as livestock grazing spread
rapidly throughout the Prairie ecoregion. Potential
conflict with agricultural interests now reduce opportunities
for species restoration in that ecoregion, and may
limit future range expansion of wolves in Montana
(Bangs et al., 1998).
A second implication of our analysis is that optimizing
both cougar and wolf populations in the same
area may be an unrealistic management goal due to interspecific
interactions. The number of wolf bounty
payments was highest where payments for cougars were
least. The spatial pattern of bounty payments indicates a
west to east gradient in abundance (least to greatest)
existed during the early 20th century for wolves, and an
opposite east to west gradient in abundance (least to
greatest) was observed for cougars. This observation is
supported by early explorer accounts of wolves in
Montana. For example, Lewis and Clark reported
wolves present throughout eastern Montana, but made
no mention of wolves once they reached the continental
divide (Coues, 1893).
Competition between wolves and cougars may exist
and have profound effects on their behavior and population
dynamics. Patterns of winter prey selection
in Glacier National Park indicated wolves and cougars
exhibit exploitation and interference competition
(Kunkel et al., 1999). Wolves can effectively compete
through direct competition for prey or occasionally
killing cougars (Bangs and Fritts, 1996; Boyd and Neale,
1992; Kunkel et al., 1999; White and Boyd, 1989). Potential
for competitive interactions is an important
conservation consideration because cougars exist in every
area where wolves are presently being restored in the
western US (Hornocker and Mobley, 1992). Cougars
are capable of exploiting valley and foothill habitats
because of relatively new, abundant white-tailed deer
(
Received 11 June 2003; received in revised form 17 October 2003; accepted 14 January 2004
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