Permeability
of the Trans-Canada Highway to Wildlife in Banff National Park: Importance of
Crossing Structures and Factors Influencing Their Effectiveness
Anthony
P. Clevenger
Department of Forestry
The University of Tennessee
Knoxville,
TN
Highway mitigation measures have been designed to
increase permeability and habitat connectivity for wildlife living in
transportation corridors. Unfortunately, post-construction performance
evaluations are rarely carried out to determine mitigation measure
effectiveness, modify them if necessary, and learn from past experiences. In
Banff National Park, Alberta, 11 wildlife underpasses (WUPs) were constructed
during the last decade to allow wildlife movement across a 4-lane section of
the Trans-Canada highway (TCH). This paper evaluates wildlife use of the WUPs
and examines the importance of structural and environmental features that may
enhance wildlife use of them. WUPs were intensively monitored during 12
months. Elk and deer used the WUPs more
frequently than carnivores. Among large carnivores, black bears were most
frequent users. Monthly crossing rates varied during the year and reflected
seasonal activity patterns. Underpass use by large carnivores was negatively
correlated with human use levels and was greatest at divided underpasses. WUP
quality was best predicted by human use levels and WUP openness and length,
yet human use was the most important factor. Ungulate use was positively
correlated with human use and negatively correlated with underpass length.
Analysis showed that human activity and openness were the most important model
components. The results suggested that possible barrier effects of the TCH may
be reduced by WUPs, but long-term studies are needed to assess TCH effects on
species' fitness. Park management of human activity around WUPs will be
crucial for success. The requirements of sensitive species should take
precedence in design of quality WUPs.
The
effect of roads on animal populations has been the focus of numerous studies
during the last few decades (Oxley et al. 1974, Muskett and Jones 1980,
Bennett 1991, Evink at al. 1996). Some well documented impacts of roads on
animal populations include: habitat loss, habitat alienation due to sensory
disturbances, barrier effects, and mortality (Adams and Geis 1983, Mansergh
and Scotts 1989, Paquet 1993, van der Zee et al, 1992). Of all these, barrier
effects or habitat fragmentation, poses what many conservation biologists
consider the greatest threat to biological diversity (Harris 1984, Saunders
and Hobbs 1991, Noss and Cooperrider 1994).
Measures
have been designed to increase permeability and habitat connectivity and
mitigate barrier effects of roads. The first wildlife crossing structures were
constructed in the 1970’s (Reed et al. 1975, SETRA 1978, Hunt et al. 1987)
and many more have been constructed since then. Nevertheless, few rigorous
evaluations regarding crossing structure effectiveness have been carried out (Romin
and Bissonette 1996).
Research aimed
at quantifying wildlife use of crossing structures (hereafter referred to as
wildlife underpasses) has focused mostly on ungulates (Reed etal. 1975, Ballon
1985, Schall et al. 1985, Singer and Doherty 1985, Woods 1990, Carsignol 1993)
and rarely on carnivores (Foster and Humphrey 1995). Only recently studies
have begun addressing what structural and environmental factors might
influence underpass use by wildlife (Foster and Humphrey 1995, Yanes et al.
1995, Rodriguez et al. 1996, Rosell et al. 1997). Virtually nothing is known
regarding differences in effectiveness between overpasses and underpasses, or
between underpass types and configurations.
Performance
evaluations of wildlife crossing structures are essential for determining
effectiveness, making recommendations for improving them if necessary, and
designing more effective measures in the future. Compared to other mitigation
measures, crossing structures, particularly those for large mammals, are
costly (Camut 1985, Gounot 1985, Leeson, this volume). Proper evaluations will
aid resource managers and highway planners when making decisions regarding the
number, placement and type of wildlife crossing structures along a section of
highway.
This paper
evaluates the patterns of wildlife use of underpasses along a 4-lane section
of the Trans-Canada highway (TCH) in the central Canadian Rocky Mountains of
Alberta. Secondly, I examine the relative importance of structural and
environmental attributes, as factors determining the quality of underpass as
perceived by wildlife.
The work was carried out in the Bow
River Valley along the TCH corridor in Banff National Park. Situated
approximately 100 km west of Calgary, BNP is the most heavily visited national
park in Canada with over 4 million visitors per year. Most of these visitors
arrive by private vehicle or motor coach along the TCH. The highway also is a
major commercial motorway between Calgary and Vancouver. Annual average daily
traffic volume at the park east entrance was 13,800 vehicles/day in 1994 and
increasing at a rate of 3% per year (R. MacMahon, Parks Canada, pers. comm.).
The transportation corridor also contains the Canadian
Pacific Railway (CPR) mainline, access roads to Banff town site and several
important two-lane highways (highways 93 and 40) and secondary roads (highway
1A). The study was conducted primarily along the easternmost 27 kilometers of
twinned (4-lane) TCH in BNP. Ten conventional highway wildlife underpasses
were constructed between 1986-88 to permit wildlife movement across the
twinned TCH. One additional underpass was constructed at Castle Junction
interchange in 1991.
A 2.4 m high
page-wire fence borders twinned sections of TCH. All underpasses are located along twinned sections of
highway. Seven of the 11 underpasses are cement open-span underpasses, two are
bridges over creeks, and two are metal culverts. Divided sections of the
highway are made up of two separate structures.
The Bow River Valley in BNP is situated within the
continental Ranges of the Southern Rocky Mountains. Elevations range from
1,300 m. to over 1,600 m. at the Continental Divide. Valley floor width varies
from 2-5 km. The climate is continental and characterized by relatively long
winters and short summers (Holland and Coen 1983). Mean annual snowfall at the
town of Banff is 249 cm. The transportation corridor traverses the Montane
Ecoregion. Vegetation consists of open forest dominated by Douglas Fir, White
Spruce, Lodgepole Pine, Aspen, and natural grasslands.
Data from
monitoring animal movements through wildlife underpasses on the TCH were used
to assess relative use of underpasses by wildlife. Wildlife visits and
through-passages at underpasses were quantified by identifying tracks of
animals on track sections at each underpass (Bider 1968). Track sections were
on level ground at underpass ends and measured roughly 2m long and 4m wide.
Tracking material consisted of a dry, loamy mix of sand. silt and clay, 3-4cm
deep. During each monitoring check, on each track section the tracks of
animals present were identified, their direction of travel recorded, and the
surface raked smooth. At each check, the tracking medium was classified as
operative or inoperative depending on the ability to read tracks clearly.
Underpasses were checked at 3-4 day intervals. Tracks of unidentified canids and small and medium-sized mammals were recorded but not included in this article. Wildlife in this study consisted of wolves, coyotes, cougars, black bears, grizzly bears, deer and elk. Other species were recorded using the underpasses (bighorn sheep, mountain goats, moose), however, due to small
sample sizes they
were excluded from the analysis. Twelve months of data were collected on
wildlife movement through the underpasses between 1 November 1996 to 31
October 1997.
To determine what
attributes most influence the use of underpass, wildlife were divided in two
guilds: large carnivores (wolves, cougars, black and grizzly bears) and
ungulates (deer and elk). Nine of the 11 underpasses were considered in the
analysis. Cascade underpass was excluded because of inconsistent and overall
sub-optimal tracking conditions. East gate underpass was left out due to its
staggered as opposed to in-line configuration thereby negating comparisons of
structural variables.
The analysis was based on through-passages by species from the two groups at the underpasses. Presence-absence data, independent of number of individuals, were used in estimating crossing rates. For each underpass, the monthly crossing index was calculated as the number of through-passes divided by the total number of through-passes at all nine underpasses during the same month. Bear
crossing data were
collected only during active months (May through October). The amount of human
activity (hikers, bikers, skiers, horses) at each underpass was quantified in
the same manner as the monthly crossing rate for wildlife.
In order to
characterize the underpasses, eight continuous variables and three categorical
variables were measured (see Table 1). Continuous variables included:
Openness (OPEN): calculated from the three former variables according to the formula (width x height/length (Reed and Ward 1985);
Three categorical variables were:
Habitat (HAB): forest (0); forest/open mix (1).
Underpass configuration (CONF): divided (0); undivided (1).
Underpass type (TYPE): open-span (0); culvert (1).
Distances between
consecutive underpass along the TCH were randomly distributed; mean distance
between underpasses was 2.7 km. indicating that crossing rates from
consecutive underpasses were independent of each other. The relationship
between the monthly crossing rate and each of the underpass variables was
examined by simple correlation (continuous variables) and non-parametric
Mann-Whitney U tests (categorical variables). Variables were log-transformed
where this provided a better fit for correlations. Forward stepwise multiple
regression analyses were then performed (Statistix 1996) to develop models
with combinations of one or more variables that explain the relative use of
underpasses. Statistical significance was set at P<0.10 in all analyses.
The underlying assumption in these analyses is that the monthly crossing rate
is a measure of the quality of that underpass as sensed by large carnivores or
ungulates.
For the 12-month
period, the average number of monthly monitoring checks at the 11 structures
was 8.5 and the average number of days between checks was 3.7 (range = 3.2 -
7.4 days). There were a total of
2,458 visits by wildlife to the underpasses (Table 2). Total number of
species' track detections at the underpasses ranged from 148 (Carrot Creek) to
482 (Buffalo). Carrot Creek had the lowest total number of animal
through-passes. The through-passage rate was highest at Buffalo, Cascade,
Edith, Powerhouse and Vermilion underpasses. Through-passage rate was lowest
at East gate (88).
There were a total
of 170 failed passages (5%), i.e., where species did not travel through the
underpasses. Monitoring checks recording no tracks occurred most often at
Morrison Coulee and Carrot Creek.
Elk were most
frequently detected at the wildlife underpasses (n-1,338, 54%), followed by
deer (n=538, 22%) and coyotes (n=373, 15%; Table 3). Among large carnivores,
black bear tracks were found 97 times (4%) at the underpasses, wolves 77 times
(3%) and cougars 29 times (1%). One wolf pack (Bow Valley pack) was
responsible for practically all the underpass use (75 out of 77
through-passes), whereas one member of the Cascade pack used the underpasses
twice during winter.
Overall
through-passage rate was high (mean = 98%, n=7), ranging from cougars and
grizzly bears (100%) to elk and deer (96%). Elk, deer, and coyotes used all of
the underpasses, while black bears were found traveling through nine, wolves
six and cougars five. Two radio-collared adult male grizzly bears used three
different underpasses.
Monthly crossing
rates for all wildlife in the study area were low from December through April,
increased sharply from May to July, and then decreased to September (Fig. 2).
There was an abrupt increase in activity during November prior to the onset of
winter. There were slight differences between large carnivore and ungulate
crossing rates over the course of the year (Fig. 3 & 4). Crossing rates
for both groups were lowest in winter. However, they differed in that large
carnivores were more active than ungulates in early spring, whereas ungulates
sustained higher crossing rates during the autumn compared to carnivores.
Correlations
between underpass variables and the monthly crossing index for wildlife
suggested that several variables were important correlates of underpass
quality (Table 4). For large carnivores, the amount of human activity was
significant and showed a strong negative correlation with underpass usage.
Underpass length was positively correlated with large carnivore crossing
indices but was not significant. Higher rates of passage were associated with
divided underpass types and were significantly different from undivided types
(P-0.07, two-tail test). All other variables showed weak correlations.
Results from
stepwise linear regression analyses of the crossing indices for large
carnivores is summarized in Table 5. Underpass quality was best predicted by
three attributes, levels of human activity, openness and underpass length,
which together explained 60% of the variance. Human activity was the most
important factor alone, accounting for more than half of the variance (30%).
Correlations
for the relationship between underpass use by ungulates and underpass
variables suggested that two variables were important in determining underpass
quality (Table 4). Level of human activity and underpass length were highly
correlated with ungulate use, the former being positive whereas the latter
negative. Ungulates were indifferent to underpass configuration, underpass
type, and type of habitat in the vicinity of an underpass failing to show any
correlations.
Underpass
quality for ungulates was best predicted by three variables, levels of human
.activity, openness and height, which combined accounted for 50 of the
variance (Table 5). Level of human activity and underpass openness were the
most important model components explaining 42% of the variation found.
Underpass length did not explain significant additional variation.
Is the
Trans-Canada highway permeable to wildlife? The results indicated that during
the 12-month period the underpasses were readily used by most species of large
carnivores and ungulates in BNP. For some species such as elk, deer, coyotes,
black bears and at least one wolf pack, crossing structures were used
regularly suggesting that possible barrier effects may be reduced. The
proportion of through-passages was high and may be a result of animals
becoming accustomed to the structures after 10 years. During a 5-year
monitoring period, (Phase 1) average through-passage rates for elk and deer
were 50% and 83%, respectively (Waters 1988). Significantly greater
through-passage rates (>96%) occurred along newly constructed underpasses
in Phase 2 (Bunyan 1990), and was best explained by prior habituation to
underpasses on Phase 1.
Monthly crossing rates from this study varied greatly
over the course of the year and were concordant with prior knowledge of
seasonal animal activity and movement patterns in the Bow Valley. Possible
reasons for the low crossing rates of large carnivores during September and
October may be related to greater or more uniformly distributed food
resources. Reduced underpass use during these months also might have resulted
from decreased bear activity prior to denning as bears accounted for half of
the large carnivores sampled.
Large carnivore
use of wildlife underpasses
was correlated with several variables; however, human use levels
was shown to be the most important and best predicted high quality
underpasses. Due to their location, several underpasses are key passages
for recreationists to cross the TCH and access popular hiking, biking
or horseback riding trails in the park. In essence, these underpasses
are conduits with elevated levels of human use and similar to
well-used trails. The influence
of humans on underpass use by large carnivores is not
surprising. In BNP and elsewhere, large carnivores generally avoid areas
of human activity including trails. Research on wolf movements in
relation to linear structures (roads, trails, railways, etc.) in the Bow
Valley showed that they tended to avoid
areas within 200m of trails (Paquet
et al 1996). Grizzly bear survival is strongly linked to areas far from
human use areas. In BNP, 100% of all grizzly bear mortality occurs
less than 500m and 200m from roads and trails respectively (Gibeau
et al 1996). Similar negative responses of black bears and cougars
to trails and human use have been reported in other areas including
national parks (Mattson et al 1987, Kasworm and Manley 1990,
McCutchen 1990, Jalkotzy and Ross 1993).
Underpass use by
ungulates was also best
predicted by the level of human
use. However, contrary to large carnivores, a strong positive correlation
was found. Most of the underpasses with high human use levels
are situated close to the town of Banff as these provide convenient
and quick access to popular hiking and horse trails The asynchronous use of
underpasses by large carnivores and ungulates can best
be explained by two separate phenomena First, large carnivores are
underutilizing or avoiding the underpasses closest to areas of significant
human activity for reasons explained above. The second is a
result of a dramatic change in elk distribution in the Bow Valley following
wolf recolonization in the early 1980's. The simultaneous recovery
of wolves and increased numbers of elk in and around the town
of Banff suggests that elk seek out wolf-free zones as refuges from predation
(Paquet et al. 1996). Deer were also considered in the analyses,
however, elk accounted for 75% of the ungulate data.
Underpass
dimensions rather than landscape attributes associated with
them have taken precedence in determining optimal structures for ungulates
(Foster and Humphrey 1995). Previous studies indicated that ungulates
were wary of using underpasses that were narrower than 7m wide
or lower than 2.4 m high (Reed et al 1975, CTGREF 1978, Velasco
et al. 1992, Rosell et al. 1997). All but one of the underpasses in
this study were greater than 7m wide (mean=9.8, SD=2.8) and all were
greater than 2.4m high (mean=3.0, SD=0.4).
Underpasses with
high openness ratios were also high quality underpasses
as perceived by ungulates. Reed et al (1975) recommended
an openness ratio for underpasses for mule deer greater than
0.6 when calculated in meters. Seven of the nine BNP underpasses surpassed this parameter (median = 0.85, range = 0.21-1.16).
Divided highway
underpasses were recommended over undivided or
bridged structures because
they are less expensive, but more importantly
the latter resemble a tunnel and may inhibit use (Foster and Humphrey
1995). The results shown here partially support that proposal
as large carnivores tended to use divided underpasses more than
undivided ones; ungulates did not select one type over the other.
The total
number of underpass visits and
through-passages was high for some species
(elk, deer, coyote), moderate for others (black bear, one wolf pack,
cougar) and extremely low for one other species (grizzly bear, one other
wolf pack). For species that used the underpasses regularly the data
indicate that possible fragmentation effects due to the TCH might be
reduced considerably. However, to determine whether the TCH (and railway)
are fragmenting wildlife populations in BNP and the Bow Valley
we need to know something about crossings and movements prior to highway construction, which unfortunately is impossible. Without
question, cross-valley and cross-TCH movements today are filtered
due to human activity, availability of quality crossing points and
underpasses along the TCH, and behavioral variability in animals As
a result, travel patterns for most animals, particularly large carnivores,
have been modified and the number of crossings are much lower
than in the past. Is this significantly affecting the fitness of animal
populations in the Bow Valley? To answer this question we will need
to carry out studies of highway effects, closely monitoring species’
movements and life history requirements in the transportation corridor.
Because many of the species of special concern are long- lived
and have low reproductive rates (e.g., grizzly bear), any deleterious
indirect effects of the highway are most likely incremental and
cumulative and many years of research will be required to assess highway
effects. The results indicate
that managing human activity around wildlife crossing structures will be of utmost importance if the structures are to
be effective in allowing wildlife to
permeate linear features such as the TCH (see Gibeau and Herrero, this volume). This has serious implications
in BNP given the current rates of growth in visitor numbers and development.
The findings described here lend greater support to the BNP management plan,
emphasizing that stricter limits to growth be imposed and calling for more
effective methods of managing and limiting human use (Banff-Bow Valley Study
1996). The permeability of the TCH has to be improved and guaranteed by a
general strategy that includes wildlife crossing structures as one measure
among others.
In addition to
restricting human use, underpasses with a large aperture should be designed in
the future. Carrot Creek, Duthil and Edith underpasses were shown to be high
quality underpasses for large carnivores and ungulates alike, primarily
because of their high openness ratio. Five-mile bridge, an unconventional
wildlife underpass along the TCH is one of the few places large carnivores
(including grizzly bears) choose to cross the TCH (Gibeau and Heuer 1996). The
bridge spans over the Bow River, the highway 1A, and the CPR mainline and is
best characterized by its great breadth and openness.
Though few habitat
or landscape attributes were measured in this analysis, none were shown to be
significant in determining quality wildlife underpasses. Nevertheless,
topography and vegetation are probably important in determining underpass
quality, and location is likely the most critical factor in guaranteeing
success. For example, large carnivores cross high-speed motorways not through
the best designed underpass, but rather through the underpass that is best
aligned with a major drainage (Beier and Loe 1992, Clevenger, unpubl. data).
In this analysis we
have attempted to address each of these factors as best possible and carry out
a rigorous evaluation of the factors most important in determining high
quality underpass for the two guilds. The results suggest that in BNP
underpasses with low human use are most important and most effective in
allowing large carnivores to permeate highways and prevent habitat
fragmentation. To design quality underpasses the requirements of sensitive
species such as large carnivores should take precedence. As such, species with
generalized habitat needs and greater adaptability to human disturbance also
will be accommodated.
Table 1
Characteristics
of nine wildlife underpasses (WUP) used in analysis of factors influencing
wildlife use.
|
WUP |
L (m) |
W (m) |
HT (m) |
OPEN |
SL (db) |
DFOR (m) |
DCPR (m) |
HUM |
HAB |
CONF |
TYPE |
|
Carrot Creek |
28.8 |
13.4 |
2.5 |
1.16 |
66.9 |
63.3 |
750 |
1.9 |
1 |
0 |
0 |
|
Morrison Coulee |
67.5
|
4.2
|
3.5 |
0.21 |
60.7 |
11.9 |
800 |
1.8 |
0 |
0 |
1 |
|
Duthil |
28.0 |
9.8 |
2.9 |
1.01 |
64.8 |
15.2 |
20 |
0.6 |
0 |
0 |
0 |
|
Power House |
27.5 |
9.5 |
2.9 |
1.00 |
64.1 |
47.3 |
20 |
5.3 |
1 |
0 |
0 |
|
Buffalo |
27.1 |
10.0 |
3.0 |
1.10 |
69.0 |
35,9 |
250 |
15.2 |
1 |
1 |
0 |
|
Vermilion |
27.2 |
9 8 |
2.7 |
.97 |
70.3 |
23.3 |
1200 |
3.2 |
0 |
1 |
0 |
|
Edith |
25.6 |
10.3 |
2.8 |
1.12 |
71.2 |
27.5 |
400 |
11.4 |
1 |
1 |
0 |
|
Healy |
28.0 |
9.0 |
2.9 |
0.65 |
66.5 |
23.9 |
750 |
0.6 |
1 |
0 |
0 |
|
Castle |
56.0 |
7.0 |
4.0 |
0.50 |
61.7 |
35.4 |
750 |
0.5 |
1 |
1 |
1 |
L;
Length, W: Width; H: Height, OPEN: openness ratio, SL: Sound level, DFOR:
distance to forest cover, DCPR: distance to CP Railroad, HU: human activity
index, HAB: habitat, CONF: configuration, TYPE: WUP type.
Table 2
Summary data from 11 wildlife underpasses (WUP) along
twinned sections of the TCH in Banff National Park, Nov. 1996-Sept. 1997.
|
WU (#
of checks) |
Total Visits |
Number
of TP’s
(%) |
Number
of failed TP’s |
No Tracks |
|
East Gate (97) |
286 |
253
(88) |
33 |
14 |
|
Carrot Creek (97) |
148 |
142
(96) |
6 |
|