Tom Paragi, ADF&G, Div. Wildlife Conservation, Fairbanks
(
During 15-18 February 2000, Al Root (ADF&G, McGrath) and I conducted browse surveys on sites along the Kuskokwim River within about 15 miles of McGrath, Alaska (Game Management Unit 19D). Previous surveys were conducted on these sites by Jack Whitman in late May or early June (1989, 1990, 1991, and 1994) in selected patches of riparian willows to assess overwinter browsing by moose. Ice scouring obliterated flagging that had marked the start and end of survey transects, so location of transects had to be surmised from 1:63,360 scale topographic maps. A scaled photograph on slide film was taken in a representative section of each transect for an archive.
Twelve surveys (see attached map for locations) were completed by sampling every five paces along 250 m transects, each a total of 50 data points, to be consistent with previous surveys. At every fifth footfall, the closest tree or shrub available to moose browsing (defined as <3.1 m [10 ft]) was categorized to species when possible. Height and distance to nearest available stem of the same species were estimated by use of a graduated meter stick. Browsing intensity class was assigned to the stem based on the count proportion of current annual growth (CAG) leaders that were browsed by moose: none, low (1-25% of leaders), moderate (26-75%), or high (76-100%). Notes were taken on the proportion of stand that had a hedged growth form (browsing intensity from previous years), visible proportion of decadent stems (largely dead), and other criteria indicative of browsing history and stand condition.
An importance index to gauge browsing intensity on a species for a given year was calculated when data for all transects were combined, similar to previous years (J. Whitman memos, 1989 and 1994, ADF&G files, McGrath). Midpoint of range for browse intensity classes (e.g., midpoint of low [1-25%] = 0.125) was multiplied by the grand mean percent frequency of samples for each browse intensity class, by species. The sum of the three products (L, M, and H classes) was then multiplied by the frequency of occurrence of the species along all transects to derive the importance index (Table 1).
Results
The vast majority of riparian willows were of the same
growth form: a stout leader up to 2 m on a single stem or small number of stems
from a common base, with CAG being smooth bark of olive or
Although the number and location of browse transects sampled has varied among years, feltleaf willow continues to be the predominant species in riparian willow stands near McGrath (Table 2). The post-winter assessments that occurred during 1989-94 showed a consistently high percentage of individual stems browsed but substantial variation in the distribution of browsing extent on those stems (importance index; Table 2). The browsing extent on feltleaf willow was higher during winters of deep snow compared to winters of normal snow depth, and in severe winters the moose sometimes browsed beyond the extent of CAG (J. Whitman, ADF&G, pers. comm.). Although the sampling in February 2000 occurred at least two months prior to the end of snow cover and beginning of new willow growth, 95% of feltleaf stems had already been browsed (Table 2), with 84% of the browsing in the high intensity class (76-100% of stems browsed per individual).
Most willow bars surveyed in past years have now grown to a height extent becoming marginally feasible for moose browsing. We observed browsing to 5 m where the moose obviously had “walked down” the stems between their front legs to reach the smaller-diameter CAG on feltleaf willow. The predictable pattern of increased height and self-thinning of stems (greater nearest-neighbor distance) over time following disturbance was reasonably evident on sites we visited (Table 3). We skipped transect 89-G (“old town slough”) because most of the plot appeared to be tall alders with very little available feltleaf or redstem willow. Three other sites near town (89-E, 89-F, and 94-H) were not sampled because of time constraints. Two of the sites were adjacent to the airport runway and had mechanical disturbance that is maintaining shorter willow. Undisturbed areas in the three sites had willows >3 m tall but were still being used by moose (A. Root, in litt.).
Overall, the hedged or broomed growth form of willows that have endured heavy, persistent browsing beyond CAG was not common in the riparian feltleaf willows we observed near McGrath. Dead stems composed <10% of areas within stands we visited, which had stem breakage only in pockets of heavy browse use. Because of relatively infrequent browsing pressure and the time lag since widespread ice scouring, substantial parts of some riparian terraces are maturing into willow forest and becoming dominated by alder thickets >4 m tall.
Discussion
Data from these browse transects and general observations
over time suggest that the intensity of browsing on riparian willows by moose
along the upper
Riparian willow stands can be critical winter forage for
moose in years of prolonged deep snowfall in this area (R. Skinner, pers.
comm.). Despite observed browsing by
moose to 5 m this year, riparian feltleaf stands on the
Methods of assessing a population size relative to its carrying capacity are challenging, both conceptually and technically. Surveys of range condition over a more inclusive cross-section of upland, lowland, and riparian willow stands would be instructive of browsing intensity and possibly predictive of population-level effects driven by nutrition. However, difficult access to large, sometimes poorly-defined seasonal ranges would require costly techniques of remote sensing and image processing, or costly access and labor for ground sampling. Rump fat or other condition indices for moose subsume the effect of range quality and weather, but they require capture and handling of individuals, which is costly and not without danger of injury or mortality to free-roaming wild animals. An annual assessment of reproductive fitness would track the response of a population to range and weather conditions without the need to capture and handle individuals. For example, neonatal twinning rate could be estimated by observing about 30 parturient cow moose from fixed-wing aircraft. Periodic range assessment, when and where feasible, would provide a complementary index to population health and possibly identify areas and objectives for habitat enhancement.
Table 1. Raw counts
and calculations for importance index of feltleaf willow in February 2000 along
the
|
Transect |
No use |
Light use |
Mod use |
Heavy use |
Total feltlf |
All spp.a |
|
89-A |
1 |
4 |
7 |
21 |
33 |
34 |
|
89-B |
1 |
0 |
4 |
36 |
41 |
50 |
|
89-D |
1 |
0 |
2 |
42 |
45 |
50 |
|
89-H |
0 |
1 |
1 |
12 |
14 |
50 |
|
94-A |
0 |
0 |
5 |
42 |
47 |
50 |
|
94-C |
2 |
0 |
3 |
41 |
46 |
50 |
|
94-G |
3 |
2 |
3 |
40 |
48 |
50 |
|
94-I |
2 |
4 |
8 |
36 |
50 |
50 |
|
94-J |
0 |
3 |
0 |
38 |
41 |
50 |
|
94-K |
1 |
3 |
6 |
36 |
46 |
50 |
|
94-M |
8 |
0 |
9 |
30 |
47 |
47 |
|
94-N |
8 |
4 |
8 |
28 |
48 |
50 |
|
|
|
|
|
|
|
|
|
Total |
27 |
21 |
56 |
402 |
506 |
581 |
|
|
|
|
|
|
|
|
|
Use freq. |
0.053 |
0.042 |
0.111 |
0.794 |
|
|
aTwo
transects were less than 50 points because at some point along transect all
willows were >3 m tall and offset continuation of the transect was not
feasible.
Proportion of all transects points that were feltleaf willow = 506/581 = 0.871
Calculation of importance index for feltleaf willow:
Browse Midpoint Feltleaf use
class of class frequency Product
Low 0.125 0.042 0.005
Moderate 0.500 0.111 0.056 Proportion
Heavy 0.875 0.794 0.695 feltleaf
-------
Sum 0.756
x 0.871 = 0.658
|
|
Table 2. Statistics
on feltleaf willow stems along the
Year #
transects % feltleaf % browsed Importance
1989 8 68 89 0.414
1990 3 39 86 0.214
1991 6 87 100 0.650
1994 14 86 85 0.307
2000 12 87 95 0.658
Table 3. Change in
mean height (m) and mean nearest-neighbor distance (m) for feltleaf willow
stems <3.1 m tall on selected transects distributed along the
Transect Metric 1989 1990 1991 1994 2000
89-A Height 2.59 -- 1.82 -- 2.07
89-A Distance 0.16 -- 0.31 -- 0.34
89-B Height 1.65 -- 1.46a -- 2.10
89-B Distance 0.41 -- 0.40a -- 0.34
89-D Height 1.11 -- -- -- 1.66 89-D Distance 0.48 -- -- -- 0.37
89-H Height 1.62 1.78 1.41b -- 2.68
89-H Distance 0.31 0.27 0.39b -- 0.59
94-G 1.91 2.49
94-G 0.52 0.44
94-I 1.30 2.45
94-I 0.34 0.92
94-J 1.63 2.28
94-J 0.30 0.48
94-K 2.11 2.29
94-K 0.26 0.45
94-M 1.10 2.28
94-M 0.18 0.38
94-N 1.86 2.08
94-N 0.22 0.38
aTwo transects were done at this site in 1991, so this is a grand mean of the two means.
bThree transects were done at this site in 1991,
so this is a grand mean of the three means.