FINAL TECHNICAL REPORT - PART A Landscape Level Hazard Rating For Western blackheaded Budworm In British Columbia

Forest Innovation Investment Number: R04-007CFS

Project Title: Development of hazard rating systems for the western hemlock looper, western blackheaded budworm and forest tent caterpillar.

Award Recipient's Name: Imre S. Otvos

Organization: Natural Resources Canada, Canadian Forest Service

Team Members: Kangakola Omendja (current term) Nicholas Conder (CFS Research Technician) Holly Armstrong (term)

Report for the Period from: April 1, 2003 to March 31, 2004 Abstract

The western blackheaded budworm (Acleris gloverana (Walshingham)) is a cyclic defoliator of western hemlock (Tsuga heterophylla (Raf.) Sarg.) and mountain hemlock (Tsuga mertensiana (Bong.) Carr.) in southern Alaska, British Columbia (BC.), and Washington State. At least 9 outbreaks of blackheaded budworm have occurred in British Columbia. Severe defoliation has been recorded repeatedly in the costal forest region of BC. Normally, the blackheaded outbreak collapses following two to four years of defoliation. To minimize the impact of defoliation by this pest, a hazard rating system has been develop to assist forest manager in dealing with outbreaks of this pest.

1 Introduction

The purpose of creating the western blackheaded budworm hazard rating system is to provide a set of tools that forest manager can utilize to increase their ability to anticipate the damaging insect activities and modify harvesting plans if necessary. The western blackheaded budworm (Acleris gloverana (Walshingham)) is a cyclic defoliator of western hemlock (Tsuga heterophylla (Raf.) Sarg.) and mountain hemlock (Tsuga mertensiana (Bong.) Carr.) in southern Alaska, British Columbia, and Washington State. Other conifer hosts, such as amabilis fir (Abies amabilis (Dougl. ex Loud.) Forbes), Douglas fir (Pseudotsuga menziesii (Mirbel) Franco), white spruce (Picea glauca (Moench) Voss), and Sitka spruce (Picea sitchensis (Bong.) Carr.), can also be attacked and defoliated when large – scale outbreaks occur in mixed stands with western hemlock (Furmiss and Carolin 1977). Periodic outbreaks of this pest result in severe defoliation in the western hemlock zones of North America (Otvos et al. 2001). At least 9 outbreaks of blackheaded budworm have occurred in British Columbia The spatial analysis of defoliation pattern of past outbreaks, and the overlay with biogeoclimatic units (Krajima 1965, Pojar et al.1987), elevation, and available climate data will help to identify and rate western hemlock stands vulnerable to western blackheaded budworm attacks. The aim of the year-end report is to identify and rate areas at risk to western blackheaded budworm outbreaks using geographical information system. Areas where outbreaks are most likely to occur can be identified and permanent pheromone trap sites can be established in these stands to monitor population changes through a complete outbreak. Monitoring trap catches will allow the detection of increasing insect defoliation (Otvos et al. 2001), which can then be used to develop a pest management system for this species similar to one developed for the Douglas–fir tussock moth (Shepherd and Otvos 1986).

2 Methods and Material

2.1 Defoliation map Annual pest surveys conducted by the Forest Insect and Disease Survey (FIDS) of the Canadian Forestry Service (CFS), form the time it was established in

2 1936 until its dissolution in 1996, used a variety of methods to determine the presence and extent of insect and disease outbreaks. Prior to the 1960s, surveys were conducted in British Columbia using boat and ground patrols, and sporadically by aircraft. From the 1960s onward, insect defoliation was mapped annually from the air, first by using fixed-wing aircraft, the later using a combination of fixed and rotary wing aircraft. Outbreaks were generally checked on the ground to verify the causal agents of the defoliation recorded from the air. Outlines of infestation were traced onto 1:250,000 scale topographic maps (NTS), producing an accurate assessment of pest activity from year to year. The historical FIDS pest outbreak survey maps were digitized and compiled into a database for the entire Province of British Columbia. The initial map projection was Universal Transverse Mercator (UTM). As coverages from each 1:250,000 NTS mapsheet were added together for each year, they were re-projected into the CFS- Pacific Region’s standard Lambert Conformal Conic Projection. For this paper, we used historic maps from all British Columbia western blackheaded budworm outbreak database from 1951 to 2001.

2.2 Data summary 1. Defoliation data: All defoliation data from 1951 to 2001 was combined together to produce a composite coverage. As data in the PFC database are in datum nad27, the composite coverage was reprojected to datum nad83 so it can be in the same datum than other data sets. 2. Biogeoclimatic zone of BC. This dataset up to date and was downloaded from the MSRM website. Initially the dataset was in BC Albers nad83 and was reprojected to Lambert nad83 to fit to the PFC official projection. 3. Elevation: the elevation coverage was interpolated from DEM point data set, using rivers and lakes as break lines.

2.3. Hardware and Software. The PFC has an extensive Geographical Information System. Most of the analysis has occurred using Environment System Research Institute’s (E.S.R.I) software program ArcGis 8.2 Desktop workstation. The data management was done within the ArcGis module ArcCatalogue, ArcTool Box, Arc and ArcEdit module. Map was produce using ArcMap module. The raster analysis was done in the GRID module. Part of the analysis was done manually or automated using AML (Arc Macro Language). Some data analysis and manipulation was done in the UNIX and Windows operating system using ArcView 3.1.

3. Defoliation pattern

3.1 Elevation In general, the majority of defoliation 90 % occurred in the elevation range of 200 – 1600 m (Fig. 1).

3 30.00

25.00

20.00

15.00

10.00

% ot Total defoliation 5.00

0.00

2200 + 200-399400-599600-799800-999 1000-11991200-13991400-15991600-17991800-19992000-2199

Figure 1: The distribution of defoliation by elevation range in meters.

The main problem of using large-scale DEM data is that it does not account for detailed topography. This can be misleading. For example, where an area of low relief in the interior of B.C. has the same elevation as high relief of the coastal forest region. The fact that the high elevation of the coastal forest region has harbored the majority of the attacks is ignored. This data set was given low weight in the final hazard rating (Fig. 2).

80

70

60

50

40

30

20 % of total defoliation 10

0 AT CWH ESSF ICH IDF MH MS SBP

Figure 2. Defoliation by biogeoclimatic zones

3.2. Biogeoclimatic Units In general most of the defoliation (73 % of total defoliation) by western blackheaded budworm occurs in the costal western hemlock biogeoclimatic zones (CWH). This biogeoclimatic zone occurs at low to middle elevations mostly west of

4 the costal mountains, along the entire British Columbia. The zone covers much of Vancouver Island, the Queen Charlotte Islands, and the Coast Mountains. The CWH is, on average, the rainiest biogeoclimatic zone in British Columbia. The mean annual temperature is about 8˚C and ranges from 5.2 to 10.5˚ C among the CWH sub-zones (B.C. Ministry of Forest, 1991). The other 27 % of defoliation occurred in decreasing order in Interior Cedar Hemlock (ICH) 15 %, Mountain Hemlock (MH) 8 %. The remaining 4 % occurred respectively within AT, ESSF, IDF, MS, SBP.

3.3. Western hemlock stands. The western hemlock is usually the most common species in the forest cover in the wet coastal region. Western hemlock regenerates freely under the canopy of mature stands on zonal sites and elsewhere if sufficient acid raw humus of decaying wood has accumulated on the forest floor (B.C. Ministry of Forest 1991). The western hemlock occurs in two ranges, a coastal range west of the Cascade and Coast Mountains, and an interior range in the Columbia Mountains west of the continental divide. (Daniel Gavin et al. 2003). These two ranges differ in term of climate and tree composition: The coastal range is generally cooler and moister than the interior. More over, the tree diversity is much greater in the interior than in the coastal range (Daniel Gavin et al.2003). In regard to tree diversity, the coastal range is likely to be more susceptible to the western blackheaded budworm defoliation because of the presence of pure hemlock stands, that is the main host for this pest.

4. Process

Rational The rational behind the development of the western blackheaded budworm hazard rating is that it is a priority to find area that bear common characteristics to regions that had experienced past WBHB outbreaks. These common conditions are mapped and added together to determine locations across the province, which shares the similar specifics parameters as the regions where WBHB occurred. This process involves the following steps:

a) Data research, gathering and manipulation b) Data mining c) Rasterization of parameters d) Proximity to past outbreaks calculation e) Weighting of parameters. f) Hazard Grid production g) Map production.

Explanation: a) The data for this specific pest is not easy to gather. The data that is in the PFC spatial database is not always complete (eg some data are missing), some data were downloaded from the forest network webpage for the missing data. Also, data were prepared and manipulated (georeferencing, attribute and spatial data integration) before analysis. b) Data mining: Use of GIS to discover the relationships among mapped variables through overlay technique (identity). All environment variables were

5 overlayed to the defoliation to determine which characteristics of environment variable are suitable for the occurrence of blackheaded budworm outbreak. c) The environment and biotic parameters was rasterize after data mining to allow of use mathematics for the production of hazard and probability surface. d) The proximity to the past outbreak was calculated through the Euclidian distance from the edge of the defoliation polygon. The distance from the polygon edge over 2 km had rate close to zero. The key distance was selected after some discussion with former FIDS rangers and specialist on how the blackheaded budworm disperses. e) Parameters were weighted following the same grid that used for the hemlock looper coarse scale hazard rating. (Otvos and Borecky 1999).

Table 1. Weighting scheme Parameter Weighing Proximity to past outbreak 20 Biogeoclimatic Zones 3 Elevation (DEM) 1.5

f) The hazard grid was produce by using map algebra and some local and focal function in the GRID module of ArcGis. Each cell value within parameters grid (proximity to outbreak, biogeoclimatic units and elevation) was calculated base on neighboring cells to take account of spatial dependency of spatial data. We use the a rectangular filter of 3 by 3 cells, of 2 km, to calculate the value of each cell base on value of surrounding cells. The resulting grids were then weighted and combine to produce the final landscape level with the cell resolution of 2 km. g) Maps were produce by the ArcMap module of ArcGIS.

5. Results and Discussion.

The final landscape level hazard rating grid has value range of 0 to 55. Each cell was then classified using one of the standard classification schemes. For western blackheaded budworm landscape level hazard rating we choose the natural breaks (jenk’s). This classification scheme finds groupings and patterns inherent in the hazard grid. So values within a class are likely to be similar, and values between classes, different. The resulting classification emphasizes the differences between classes (high, medium, low and minimal). The data was classified as follows: < 5 minimal hazard, 5 – 15 low hazard, 15 – 33 medium hazards and > 33 high hazard. Figure 3 gives an overview of the areas susceptible to western blackheaded defoliation in British Columbia. The inset area detailed in the Coastal forest region around Queen Charlotte Islands. The highest susceptible areas conform to location where defoliation has occurred in the past. For this hazard rating, proximity to past defoliation areas and biogeoclimatic are the core data set. Area that has suffered by defoliation in the past is likely to be defoliated again in the future. More over, as reported by Otvos et al. (2000), the biogeoclimatic zone is a much better indicator of forest stand susceptibility to defoliation than forest stand characteristics (density,

6 proportion of hemlock, air crown closure and site index). A limitation with the present hazard-rating mapping is the unavailability of accurate location of hemlock mature stand. This current hazard rating at landscape level will be improved by incorporating a more accurate information on the mature stand location in the hazard rating when those become available.

Figure 3. Western blackheaded budworm Landscape Level Hazard Rating.

The present hazard mapping provides decision support when considering the placement of pheromone trapping to monitor the evolution of western blackheaded budworm.

6. Conclusion.

In future the effort should be concentrated on the refinement of the landscape level hazard rating by incorporating more accurate information on the location of mature hemlock stands (as they become available) and the creation of stand level hazard rating and risk assessment. This will help us to rate hemlock stand and

7 calculate the stand risk to defoliation by western blackheaded budworm. To verify our current hazard rating it will be desirable to use another method, i.e. logistic regression and weight of evidence probability to create the defoliation probability surface for future western blackheaded budworm outbreak. This will be useful to forest manager for the identification of areas at risk for defoliation and use it as an input in their forest management strategies.

A preliminary hazard rating system, this is planned for publication within the next 12 months (see Appendix I)

References Borecky, N., and Otvos, I.S. 2001. Coarse-scale hazard rating of western hemlock looper in British Columbia. In Proceedings: Integrated Management and Dynamics of Forest Defoliating Insects. Victoria, British Columbia, Canada. 15- 19 Aug., 1999. Edited by A.M. Liebhold, M.L McManus, I.S. Otvos, and S.L.C. Fosbroke. USDA For. Serv. Gen. Tech. Rpt. NE-277. pp. 6-15.

Gavin, D.G., and Hu, F.S. 2003. Climatic vs. non-climatic control of western hemlock distribution in its coastal and interior ranges. Paper presented at the 2003 Annual Meeting of the Ecological Society of America, August 3-8, 2003, Savannah, Georgia. www.life.uiuc.edu/hu/gavin/pubs.html

Furniss, R.L. and Carolin, V.M. 1977. Western Forest Insects. USDA Forest Service Misc. Publ. 1339. USDA Forest Service, Washington, DC. Pp. 205-208.

Krajima, V.J. 1965. Biogeoclimatic zones and classification of British Columbia. Ecology of Western North America 1: 1-17.

Meidinger, D., and Pojar, J. 1991. Ecosystems of British Columbia. British Columbia Ministry of Forests Special Report Series 6. Research Branch, British Columbia Ministry of Forests, Victoria, B.C.

Otvos, I.S., and Borecky, N. 1999. Western Hemlock Looper Hazard Rating System WHLHRS: Preliminary Report. Unpublished Report.

Otvos, I.S., Borecky, N., Shepherd, R.F., and Dewey, A. 2001. Spatial relationships between western blackheaded budworm (Acleris gloverana) (Lepidoptera: Tortricidae) defoliation patterns and habitat zones on Vancouver Island, British Columbia. In Proceedings: Integrated Management and Dynamics of Forest Defoliating Insects. Victoria, British Columbia, Canada. 15-19 Aug., 1999. Edited by A.M. Liebhold, M.L McManus, I.S. Otvos, and S.L.C. Fosbroke. USDA For. Serv. Gen. Tech. Rpt. NE-277. pp. 133-143.

Pojar, J., Klinka, K., and Meidinger, D.V. 1987. Biogeoclimatic ecosystem classification in British Columbia. For. Ecol. Manage. 22: 119-154.

Shepherd, R.F., and Otvos, I.S. 1986. Pest management of Douglas-fir tussock moth: procedures for insect monitoring problem evaluation and control actions. Pac. For. Centre, Inf. Rpt. BC-X-270.

8 Appendix I

Title of Proposed Scientific Paper to be Published Within the Next 12 Months

Development of a Western Blackheaded Budworm, Acleris gloverana (Lepidopter: Tortricidae), Hazard Rating System and Risk Probability Surface Using Logistic Regression and Bayes’ Theorem

By

Kangakola Omendja and Imre S. Otvos

9 700,000 Outbreak 1 600,000

500,000

400,000 Outbreak 5

300,000 Area defoliated (ha) 200,000 Outbreak 2

Outbreak 7 Outbreak 9 100,000 Outbreak 3 Outbreak 4 Outbreak 6 Outbreak 8

0

1951 1954 1957 1960 1963 1966 1969 1972 1975 1978 1981 1984 1987 1990 1993 1996 1999

Figure 1. Total area of defoliation (light, moderate and severe) by western blackheaded budworm, as recorded by the Canadian Forest Services Forest Insect and Disease Survey (from 1951 to 2001).

10 1200000 73.5 %

1000000

800000

600000

22.1 % 400000 Area defoliated (ha)

200000 < 1 % 3.6 % < 1 % < 1 % < 1 % 0 1 2 3 4 5 6 7 Total number of years

Figure 2. Cumulative years of defoliation of the same area by the western blackheaded budworm in British Columbia, 1951-2001.

11 300000

250000

High (>5 years)

200000 Medium (3-4 years) Low (1-2 years)

150000 Area defoliated (ha) 100000

50000

0

AT un ICH dm IDF dw SBS dk ESSFmc ICH vk 1 ESSFvcp ESSFwv ICH mc ICH2 mw 2 MH mm MH2 wh 1MS dc 2 CWH ds CWH1 mmCWH 2 vh CWH1 vmCWH 1 whCWH 1 ws CWH1 xm 2 ESSFwc ESSFwc2 ESSFwcw5 Biogeoclimatic zone

Figure 3. Susceptibility of hemlock stands to western blackheaded budworm in defoliated biogeoclimatic zones based on western blackheaded damage from 1951 to 2001.

NB This figure is intended for formal journal publication (see Appendix I for title).

12 Table 1. Total area of defoliation (ha) recorded for each Forest Region in British Columbia from 1951 to 2001 Coast Northern TOT YEAR al Interior Southern Interior Outbreak AL

1951 1673 1673 1952 18660 3378 22037 1953 184674 234001 418675 1954 307541 293788 601329 Subtotal outbreak 1 1043715 1959 50629 50629 1960 107613 17931 125544 Subtotal outbreak 2 176174 1965 2617 2617 1966 76635 76635 Subtotal outbreak 3 79252 1968 458 458 1969 10610 10610 Subtotal outbreak 4 11068 1971 48571 48571 1972 164007 164007 1973 34954 31872 66825 1974 123632 171339 294971 Subtotal outbreak 5 574374 1976 9025 9025 1977 95 95 Subtotal outbreak 6 9121 1981 100 100 1983 229 229 1984 21803 21803 1985 77542 951 4767 83260 1986 58443 58443 1987 2213 2213 1988 4898 1766 6664 1989 11956 11956 1990 856 856 Subtotal outbreak 7 185524 1993 46 46 1994 6041 6041 Subtotal outbreak 8 6087 1996 9433 9433 1997 10686 10686 1998 74437 74437 1999 55175 55175 2000 31293 31293 2001 1845 1845 Subtotal outbreak 9 182869

Total 1376869 754933 136382 2268185 % 61 33 6 100

13 Table 2 Western blackheaded budworm-defoliated hemlock stands by biogeoclimatic zones during all previously recorded outbreaks

Area % of total % of Biogeoclimatic defoliated BGCZ area total area variant Description (ha) defoliated defoliated AT un Alpine Tundra 3301.73 0.09 0.22 AT unp Alpine Tundra 8612.18 0.08 0.57 Total Alpine Tundra 11913.91 0.79

CWH dm Coastal Western Hemlock dry mild 165.60 0.03 0.01 Coastal Western Hemlock dry submaritime CWH ds 1 variant 1 64.85 0.02 0 Coastal Western Hemlock dry submaritime CWH ds 2 variant 2 828.93 1.02 0.05 Coastal Western Hemlock moist maritime CWH mm 1 variant 1 5970.33 3.95 0.39 Coastal Western Hemlock moist maritime CWH mm 2 variant 2 22437.64 9.80 1.48 Coastal Western Hemlock moist maritime CWH ms 1 variant 1 395.83 0.08 0.03 Coastal Western Hemlock moist maritime CWH ms 2 variant 2 2279.42 1.71 0.15 Coastal Western Hemlock very wet CWH vh 1 hypermaritime variant 1 3268.30 0.69 0.22 Coastal Western Hemlock very wet CWH vh 2 hypermaritime variant 2 196587.58 11.56 12.99 CWH vm Coastal Western Hemlock very wet maritime 90851.31 30.83 6 Coastal Western Hemlock very wet maritime CWH vm 1 variant 1 181565.01 8.60 12 Coastal Western Hemlock very wet maritime CWH vm 2 variant 2 103149.73 7.42 6.82 Coastal Western Hemlock very wet maritime CWH vm 3 variant 3 330.54 0.44 0.02 Coastal Western Hemlock wet hypermaritime CWH wh 1 variant 1 213175.11 38.14 14.09 Coastal Western Hemlock wet hypermaritime CWH wh 2 variant 2 31067.14 37.06 2.05 CWH wm Coastal Western Hemlock wet maritime 41775.50 12.49 2.76 Coastal Western Hemlock wet submaritime CWH ws 1 variant 1 138139.61 55.92 9.13 Coastal Western Hemlock wet submaritime CWH ws 2 variant 2 68264.29 10.34 4.51 Coastal Western Hemlock very dry maritme CWH xm 1 variant 1 17.00 0.00 0 Coastal Western Hemlock very dry maritme CWH xm 2 variant 2 3777.02 0.75 0.25 Total Coastal Western Hemlock 1104110.74 72.97

14 Area % of total % of Biogeoclimatic defoliated BGCZ area total area variant Description (ha) defoliated defoliated Engelmann Spruce-Subalpine Fir dry cold variant ESSFdc 2 2 353.55 0.15 0.02 Engelmann Spruce-Subalpine Fir dry mild variant ESSFdm 1 1 94.58 0.06 0.01 ESSFmc Engelmann Spruce-Subalpine Fir moist cool 2895.57 0.26 0.19 ESSFmcp Engelmann Spruce-Subalpine moist cool p. 267.76 0.13 0.02 ESSFvc Engelmann Spruce-Subalpine Fir very wet cold 7443.11 2.20 0.49 ESSFvcp Engelmann Spruce-Subalpine Fir very wet cold p. 18.43 0.01 0 ESSFvv Engelmann Spruce-Subalpine Fir wet 19.99 0.02 0 Engelmann Spruce-Subalpine Fir wet cold variant ESSFwc 1 1 4211.82 1.90 0.28 Engelmann Spruce-Subalpine Fir wet cold variant ESSFwc 2 2 5186.06 0.61 0.34 Engelmann Spruce-Subalpine Fir wet cold variant ESSFwc 3 3 514.79 0.06 0.03 Engelmann Spruce-Subalpine Fir wet cold variant ESSFwc 4 4 4662.28 0.80 0.31 Engelmann Spruce-Subalpine Fir wet cold variant ESSFwc 5 5 134.58 0.73 0.01 Engelmann Spruce-Subalpine Fir wet cold variant ESSFwc 6 6 4.18 0.01 0 ESSFwcp Engelmann Spruce-Subalpine Fir wet cold p. 594.19 0.10 0.04 ESSFwcw Engelmann Spruce-Subalpine Fir wet cold w. 8.44 0.01 0 Engelmann Spruce-Subilpiene Fir dry cold wet ESSFwk 1 cool variant 1 2326.19 0.43 0.15 ESSFwm Engelmann Spruce-Subalpine Fir wet mild 179.05 0.06 0.01 ESSFwv Engelmann Spruce-Subalpine Fir wet 675.32 0.04 0.04 ESSFxv 1 Engelmann Spruce-Subalpine very dry very cold 2079.16 0.66 0.14 Total Engelmann Spruce-Subalpine Fir 31669.05 2.09

ICH dw 1 Interior Cedar Hemlock dry warm variant 1 298.61 0.07 0.02 ICH mc 1 Interior Cedar Hemlock moist cold variant 1 38811.31 7.38 2.56 ICH mc 2 Interior Cedar Hemlock moist cold variant 2 82806.97 25.36 5.47 ICH mk 1 Interior Cedar Hemlock moist cool variant 1 39.22 0.01 0 ICH mw 1 Interior Cedar Hemlock moist warm variant 1 163.49 0.11 0.01 ICH mw 2 Interior Cedar Hemlock moist warm variant 2 31885.74 3.63 2.11 ICH mw 3 Interior Cedar Hemlock moist warm variant 3 2244.56 0.46 0.15 ICH mw 4 Interior Cedar Hemlock moist warm variant 4 226.88 0.48 0.01 ICH vk 1 Interior Cedar Hemlock very wet cool variant 1 18135.31 7.29 1.2 ICH wk 1 Interior Cedar Hemlock wet cool variant 1 42271.93 7.28 2.79 ICH wk 2 Interior Cedar Hemlock wet cool variant 2 2537.39 1.48 0.17 Total Interior Cedar Hemlock 219421.41 14.5

IDF dw Interior Douglas-fir 1120.97 1.06 0.07 IDF ww Interior Douglas-fir wet warm 30.40 0.02 0 Total Interior Douglas-fir 1151.37 0.08

15 Area % of total % of Biogeoclimatic defoliated BGCZ area total area variant Description (ha) defoliated defoliated MH mm 1 Mountain Hemlock moist mild variant 1 55041.19 3.40 3.64 MH mm 2 Mountain Hemlock moist mild variant 2 24881.27 2.03 1.64 MH mmp Mountain Hemlock moist mild p. 11714.28 5.56 0.77 MH un Mountain Hemlock un 1476.95 0.46 0.1 MH wh 1 Mountain Hemlock wet hot variant 1 14102.54 11.73 0.93 MH wh 2 Mountain Hemlock wet hot variant 2 4929.71 25.93 0.33 MH whp Mountain Hemlock wet hot p. 2727.75 9.11 0.18 Total Mountain Hemlock 114873.69 7.59

MS dc 2 Montane Spruce dry cold variant 2 2674.94 6.08 0.18 MS xv Montane Spruce very dry very cold 1650.60 0.18 0.11 Total Montane Spruce 4325.54 0.29

SBPSxc Sub-Boreal Pine -Spruce very dry cold 2382.32 0.22 0.16 Total Sub-Boreal Pine 2382.32 0.16

SBS dk Sub-Boreal Spruce dry cool 3542.76 0.29 0.23 SBS mc 2 Sub-Boreal Spruce moist cold variant 2 19782.10 0.91 1.31 Total Sub-Boreal Spruce 28089.50 1.54

16