Canadian Journal of Forest Research

Lifeboat or sinking ship: Will the size and shape of Old- Growth Management Areas provide viable future habitat for temperate rainforest lichens?

Journal: Canadian Journal of Forest Research

Manuscript ID cjfr-2019-0381.R2

Manuscript Type: Article

Date Submitted by the 24-Mar-2020 Author:

Complete List of Authors: Bezzola, Aita; University of Northern British Columbia, Ecosystem Science and Management Coxson, Darwyn; Univ No British Columbia, Ecosystem Science and ManagementDraft managed forests, protected areas, lichens, temperate rainforest, edge Keyword: effects

Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? :

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1 Lifeboat or sinking ship: Will the size and shape of Old-Growth Management Areas provide

2 viable future habitat for temperate rainforest lichens?

3

4

5 Aita Bezzola and Darwyn Coxson1.

6

7 Ecosystem Science and Management Program

8 University of Northern British Columbia

9 3333 University Way, Prince George, B.C., V2N 4Z9

10

11 1. Corresponding author (email: [email protected] , phone: 250-960-6646)

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12 Abstract:

13 The Kispiox Timber Supply Area, a 1.3 million ha region in northwestern British Columbia,

14 supports a significant assemblage of old-forest dependant temperate rainforest (oceanic) lichens.

15 Given their known sensitivity to , we ask whether or not the current configuration of

16 Kispiox Old-Growth Management Area reserves (OGMAs) will provide viable future habitat for

17 oceanic lichens as surrounding landscapes are progressively logged in coming decades.

18 Landscape indicators were calculated from provincial map datasets. Old cedar-hemlock forests,

19 the primary habitat for Kispiox oceanic lichens, had a landscape shape index of 6.4 in OGMAs,

20 indicative of elongate shapes susceptible to edge effects. Mean patch size in OGMAs was 43 ha,

21 with the largest patch size 1,378 ha. In contrast, the landscape shape index for pre-industrial old

22 cedar-hemlock forests was 1.3, with a meanDraft patch size of 1293 ha, and largest patch size 23,357

23 ha. When modelled edge effects were extended to 120 m, only 25% of cedar-hemlock forests in

24 Kispiox OGMAs remained interior habitat (7,754 ha total). Adoption of silvicultural practices

25 that maintain buffer zones around existing OGMAs, and the designation of additional OGMAs,

26 especially in watersheds with intact old cedar-hemlock forests, is recommended to conserve

27 oceanic lichen communities in the Kispiox.

28

29

30 Keywords: managed forests, protected areas, edge effects, lichens, temperate rainforest, variable-

31 retention harvesting

32

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33 Introduction

34 Wet-temperate rainforests in the headwaters of the Skeena and Nass River watersheds in

35 northwestern British Columbia (B.C.) represent a hotspot for old-forest dependant

36 temperate rainforest (oceanic) lichen species in western North America (Goward 1994; Goward

37 and Spribille 2005). This area falls within a zone of ecological transition, between the very-wet

38 coastal mountain ranges, where canopy lichens are outcompeted by mosses and liverworts, and

39 drier central-interior plateau habitats, where summer is insufficient to support

40 temperate rainforest lichen communities (Goward and Spribille 2005). This ecosystem was

41 recently described as the Kispiox Inland Temperate Rainforest (Coxson et al. 2019). 42 As a group, canopy lichens haveDraft commonly been used as indicators, both for their 43 sensitivity to air pollution, and as markers of forest successional stages (Esseen and Coxson

44 2015). Conditions suitable for the establishment and growth of old-forest dependant oceanic

45 lichen communities typically do not occur for upwards of two centuries or more after stand-

46 replacing disturbance events (Campbell and Fredeen 2004; Price et al. 2017). This is a reflection

47 of the time needed for development of the stand structural complexity required by habitat

48 specialists (Goward 1994).

49 Over the next five to six decades, most timber supply areas in B.C. will see a transition

50 from relying mainly upon old-forests as their primary timber supply to using second-growth

51 stands to meet timber supply needs, a transition described as the “fall-down” effect (Marchak et

52 al. 1999). Within the Kispiox region, the timber supply from natural (predominantly old) cedar-

53 hemlock forests is expected to fall by 75% or more within the next 50 years (Timberline 2007;

54 Snetsinger 2008). Habitat for old-forest dependant oceanic lichens in these future landscapes will

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55 consequently be highly constrained, with current forest management decisions having a major

56 influence on the long-term viability of oceanic lichen communities in the Kispiox.

57 Although lichen communities can be conserved on retained individual (green) trees

58 within regenerating cutblocks in managed forests (Hofmeister et al. 2016), this mainly benefits

59 species that already grow in open light environments (Lundstrom et al. 2013). For old-forest

60 dependant lichens, the post-harvest transition to high light and desiccating environments

61 typically leads to widespread dieback of canopy lichen communities (Gauslaa and Solhaug

62 1996). Old-forest dependant lichens with a cyanobacterial photosynthetic partner (symbiont), in

63 particular, are highly sensitive to prolonged high light exposure when dry (Gauslaa et al. 2006).

64 Creating suitable spatially-defined reserves where old-forest conditions are maintained is

65 therefore generally regarded as a preferredDraft conservation strategy for oceanic lichen species

66 (Ylisirniöa and Hallikainen 2018).

67 The concept of designating forestry retention patches or reserves within the timber

68 harvesting landbase to act as refugia or “lifeboats” for old-forest dependant biota such as lichens

69 is well established in forest management practices across North America and Europe

70 (Matveinen-Huju et al. 2006; Gustafsson et al. 2010; Gustafsson et al. 2012; Mori and Kitagawa

71 2014). Perhans et al. (2009), for instance, describes how forest reserves function as “lifeboats”

72 for old-forest dependant lichens during the forest regeneration phase, though cautioning that

73 small retained forest fragments or reserves may be unable to perform this function. Three major

74 factors have repeatedly been raised when the design of forest reserves in managed landscapes is

75 discussed.

76 Reserve placement must first consider the adjacency of other reserves. Old-forest

77 dependant lichens commonly have limited dispersal abilities (Sillett et al 2000; Öckinger et al.

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78 2005). Aune et al. (2005) raised concerns about the increasing isolation of forest reserves in

79 Sweden, recommending both the designation of larger reserves and the increased utilization of

80 buffer zones around reserves so that matrix landscapes can better support dispersal of old-forest

81 dependant taxa between reserves.

82 Second, reserve design must consider the sensitivity of old-forest dependant lichens to

83 edge effects, especially when considering the size and shape of forest reserves. This concern

84 arises from the requirement of many old-forest dependant lichens for ‘interior” forest conditions,

85 due to their intolerance of extremes of desiccation, high light exposure, and thermal conditions

86 (Esseen and Renhorn 1998; Gehlhausen et al. 2000; Hilmo and Holien 2002; Gauslaa et al.

87 2006). Cantú-Salazar and Gaston (2010), in discussing the relative contributions of many small

88 versus few large protected areas, note thatDraft the high perimeter-to-area ratios of small reserves

89 frequently limits their effectiveness, allowing external conditions to penetrate reserve

90 boundaries.

91 Third, habitat quality within designated reserves is critical to sustaining lichen

92 communities. Radies et al (2009) found that the distribution and abundance of old-forest

93 dependant temperate rainforest lichens was greatest in wet nutrient-receiving sites (wet “toe-

94 slope” positions), sites that historically had long time intervals between stand destroying events

95 such as fire. Fritz et al. (2008) found that long-term site continuity (>350 years) was a critical

96 factor in predicting the presence of red-listed lichens in retained forest patches in managed

97 landscapes of southern Sweden, noting that the generally higher quality habitat in patches with

98 long-term site continuity was an important contributing factor. Franklin et al. (2007) emphasized

99 the importance of providing continuity of forest cover in reserves, including both compositional

100 (tree species diversity) and structural (such as snags and coarse woody debris) elements. These

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101 considerations are particularly important in temperate rainforest ecosystems of western North

102 America, where prevailing natural disturbance dynamics historically allowed the development of

103 landscapes dominated by old-forests (Beese et al. 2003; DeLong 2007; DellaSala et al. 2011;

104 Stevenson et al. 2011; Alaback et al. 2013).

105 One of the major ways in which protection of old-forests is achieved within the timber

106 harvesting land base in B.C. is by the designation of Old-Growth Management Areas (OGMAs),

107 typically established as spatially defined reserves for old-forest protection (Environmental Law

108 Centre 2013). This stands in contrast to aspatial old-forest management targets, which set

109 landscape-level thresholds for old-forest retention but do not require designated reserve

110 boundaries (British Columbia Forest Practices Code 2004; British Columbia Forest Practices

111 Board 2012). The importance of spatiallyDraft defined conservation reserves in landscapes managed

112 for timber production has repeatedly been noted by conservation biologists, who emphasize that

113 providing protection in national and/or provincial and state parks alone cannot meet landscape-

114 level biodiversity objectives (Hansen et al. 1991; Stevenson et al. 2011).

115 Given the importance of the Kispiox to lichen conservation, a key question for

116 conservation biologists is whether or not currently designated OGMAs will function as a viable

117 refugia or “lifeboats” for temperate rainforest lichens, especially over the next 50-60 years’ as

118 surrounding landscapes are progressively harvested (Timberline 2007). To examine the future

119 viability of Kispiox OGMAs for old-forest dependant lichens, we have asked the following

120 questions: 1) what is the size class distribution and adjacency of existing OGMAs? 2) what

121 proportion of OGMAs will remain as interior-forest habitats if adjacent forests are clearcut,

122 taking into account modelled edge effects? and 3) what risks do current patterns of harvesting

123 pose to lichens in Kispiox OGMAs? The answers to these questions are placed in context of

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124 historical patterns of old-forest distribution and the location high-quality lichen habitats in the

125 Kispiox. Based on these findings we consider whether or not currently designated Kispiox

126 OGMAs will function as “lifeboats” for old-forest dependant lichens, allowing viable

127 populations to persist as surrounding landscapes are progressively logged, or whether they

128 should instead be regarded as “sinking ships”, unable to retain their lichen populations over time.

129 We also examine potential strategies that Kispiox forest managers could adopt to increase the

130 viability of Old-Growth Management Areas as refugia for old-forest dependant lichens.

131

132 Methods

133 Draft

134 Study Area

135 The Kispiox Timber Supply Area covers 1,301,220 ha within the watershed of the Skeena River

136 in northwestern British Columbia and is the primary administrative unit for making decisions on

137 timber supply allocation in this region (British Columbia Ministry of Forests and Range 2008).

138 We refer to this area collectively as the Kispiox (Fig. 1). Lands within the Kispiox fall within

139 the traditional territories of the Gitksan and Tsimshian First Nations (Suttles and Sturtevant

140 1990).

141 Our study area was comprised of old (age class 8 – forests 141-250 years in age) and

142 very-old (age class 9 – forests > 250 years in age) Interior Cedar-Hemlock (ICH) biogeoclimatic

143 zone forests in the Kispiox (Fig. 1), including the Nass Moist Cold and Hazelton Moist Cold

144 biogeoclimatic subzones (Meidinger and Pojar 1991), based on subzone boundaries and age class

145 mapping from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and

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146 Rural Development (2018a). We refer to these old and very-old ICH forest stands as old cedar-

147 hemlock forests in our present discussion.

148 Kispiox forests are dominated by western hemlock (Tsuga heterophylla (Raf.) Sarg.),

149 with a mix of other secondary trees species, including western redcedar (Thuja plicata Donn ex

150 D. Don in Lamb.), subalpine fir (Abies lasiocarpa (Hook.) Nutt.), lodgepole pine (Pinus contorta

151 var. latifolia Engelm.), hybrid spruce (the complex of white spruce (Picea glauca (Moench)

152 Voss), Sitka spruce (Picea sitchensis (Bong.) Carr.), paper birch (Betula papyrifera Marsh.),

153 trembling aspen (Populus tremuloides Michx.), and black cottonwood (Populus balsamifera ssp.

154 Trichocarpa Torr. & Gray) found as lessor components within stands (Coates et al. 1997). 155 The network of forest access roadsDraft in the Kispiox extends into valley-bottom locations in 156 all but the northern third of the region (Fig. 1), providing a close approximation of areas where

157 clearcut harvesting blocks have been established by forest industry licensees. A detailed map of

158 the road network and forest harvesting blocks in the central-Kispiox area is shown in Fig. 2.

159 Three major provincial parks are found within the Kispiox: Swan Lake Kispiox River

160 Provincial Park in the northwest (62,255 ha), Babine River Corridor Provincial Park (15,359 ha)

161 in the east, and Babine Mountains Provincial Park (31,146 ha) in the south (Fig. 1). Of these

162 three parks, Swan Lake Kispiox River Provincial Park contains the largest amount of old cedar-

163 hemlock forest habitat (Fig. 1). The land base within these parks has been excluded from our

164 analysis, which was confined to the timber harvesting land base.

165

166 Edge effects in old-forest dependant lichens

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167 The Kispiox region is home to a rich suite of temperate rainforest or oceanic lichens

168 (Coxson et al. 2019). The abundance and distribution of the smoker’s lung lichen (Lobaria

169 retigera (Bory) Trevisan) is characteristic of many of these species, occurring in both coastal and

170 inland mountain ranges (Fig. 3A). Notwithstanding its large geographic range, both habitat

171 modelling and known occurrences suggest that the vast majority of thalli are confined to the

172 Kispiox (Fig. 3B). Smoker’s lung is also a species for which we have good evidence for

173 sensitivity to edge effects. Gauslaa et al. (2018) found that thalli were severely bleached (i.e. the

174 algal photobiont becomes incapable of photosynthesis) 80 m from clearcut edges, with edge

175 effects still noticeable 120 m from edges (Gausla et al. 2018). Notably, these measurements were

176 taken on east-facing clearcut edges in the Kispiox, thus edge effects may be even more severe on

177 south- and/or west-facing aspects. ConsistentDraft with these findings, we have used distances from

178 edge of 35, 80, and 120 m to calculate scenarios for the amount of interior forest habitat

179 available for oceanic cyanolichens in Kispiox OGMAs. Our 80 and 120 m thresholds provide

180 biologically meaningful edge effect thresholds, while the 35 m threshold applied is closer to

181 current assumptions made by forest licensees when assessing edge versus interior habitat (e.g.

182 see CANFOR 2006).

183

184 Landscape metrics

185 The calculation of landscape metrics for OGMAs within the Kispiox (see Fig. 4 for

186 location of OGMAs) followed analytical methods of Armenteras et al. (2003). Indices calculated

187 were patch number, largest patch index, mean patch size, mean nearest neighbour distance,

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188 landscape shape index, total patch size, and the largest patch size1. Our patch analysis on

189 OGMAs within the Kispiox was conducted at several geographic scales. From largest to

190 smallest, these were: 1) All OGMAs containing old cedar-hemlock forest habitats (age classes 8

191 and 9), with individual OGMAs treated as discrete patches. This analysis is comparable to

192 existing ministry databases describing the total number of OGMAs and their area. It should be

193 noted, however, that many of these OGMAs are comprised predominantly of other forest types

194 (with cedar-hemlock a minor component), so this analysis is limited in its ability to predict

195 conservation values for oceanic lichens, which occur mainly in old cedar-hemlock forests. 2) Our

196 primary analysis focus was therefore on the size and pattern of the actual old cedar-hemlock

197 forest patches (age classes 8 and 9) within OGMAs, with each discrete area of cedar-hemlock

198 forest treated as an individual patch. ThisDraft analysis provides the most meaningful prediction of

199 current and future habitat availability for oceanic lichens in the Kispiox, confined as it is to the

200 actual habitat for these lichens. Our subsequent analysis examines how much of this cedar-

201 hemlock forest habitat in OGMAs will remain viable habitat after the impacts of edge effects are

202 considered. These analyses were: 3) As in (2), but including only areas of “interior habitat”

203 forest that were more than 35 m from edges of OGMAs; 4) As in (2), but including only areas of

204 ‘interior habitat” that were more than 80 m from edges of OGMAs; 5) As in (2), but including

205 only areas of “interior habitat” that were more than 120 m from edges of OGMAs. The patch size

206 distribution (number of patches) for discrete cedar-hemlock forest patches (categories [2] to [5]

207 above) were categorized for patch sizes of < 1, 1-5, 5-10, 10-20, 20-50, 50-100, and 100+ ha.

208 Our analysis includes previously established OGMAs within the eastern part of the

209 Kispiox (in the Xsu gwin lik"l"inswx: West Babine Sustainable Resource Management Area)

1 Please refer to supplemental materials F and G for a more detailed description of methodologies and data sources for landscape metrics. Page 10

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210 that have now been designated as Core Ecosystem reserves (British Columbia Ministry of

211 Forests, Lands, Natural Resource Operations and Rural Development 2019), excluding any

212 reserves subsequently incorporated into the Babine River Corridor Provincial Park. For the

213 purposes of the present discussion we will refer to both of these reserve types (OGMAs and Core

214 Ecosystem reserves) as OGMAs; each represents spatially designated areas within the

215 boundaries of the timber harvesting land base that are excluded from forest harvesting.

216 The same landscape metrics were calculated for old cedar-hemlock forest stands in

217 “matrix” (surrounding) forests within the larger landscape. These calculations were made for two

218 different sets of study areas within the Kispiox: first within a representative 27,819 ha area of

219 roaded (prior first-pass clearcut logging initiated) old cedar-hemlock forest landscape in the

220 central-Kispiox (Fig. 1A); and second, withinDraft four remaining “unlogged” (unroaded) watersheds

221 with a total area of 30,859 ha (Figs. 1B – Kuldo Creek, 1C - Upper Skeena, 1D – Sicintine River,

222 and 1D - Shedin Creek). Our analysis on the central-Kispiox additionally included calculation of

223 landscape metrics for what the state of “preharvest” forests in this area would have looked like.

224 This extrapolation to “preharvest” conditions assumed that currently mapped cutblocks (since

225 1980) had been harvested in age class 8 or 9 forest stands. We additionally removed roads from

226 the “preharvest” analysis. This analysis will slightly underestimate the amount of old-forests in

227 the “preharvest” landscape as we do not account for older cutblocks (which are treated as

228 younger age class stands in our analysis) or for other disturbances types such as fire (which are

229 rare in this landscape).

230 Geographic analysis and mapping was done using ArcMap release 10.5.1 (ESRI,

231 Redlands, CA. http://desktop.arcgis.com/en/arcmap/) and QGIS release 3.2, an Open Source

232 Geographic Information System (https://qgis.org/en/site/).

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233 Several measures were taken to ensure that our edge effect and shape parameter

234 calculations were not unduly influenced by artifacts in the mapping data. First, where OGMAs

235 contained small patches within their boundaries that had been excluded from formal designation

236 as OGMAs (e.g. small within OGMAs), but were surrounded on all sides by OGMAs,

237 these patches were treated as if they were a part of OGMAs. Therefore, natural “internal” edges

238 were generally excluded from our analysis, which focused on “outer” edges. Second, all mapped

239 OGMAs that were smaller than 1 ha in size were removed from the dataset, removing possible

240 artifacts created by mapping irregularities. We would note that the dataset contained many

241 fragments (often referred to as slivers) that seem to be artifacts of the overlay and rectification of

242 different map layers. These are unlikely to be part of bona-fide OGMAs, rather they likely

243 represent residuals from spatial data processing.Draft Cumulatively, these fragments represented

244 0.65% of the database layer for OGMAs.

245 The B.C. Vegetation Resources Inventory dataset was subsequently reduced to subzone

246 boundaries for the Interior Cedar-Hemlock biogeoclimatic zone, which were then clipped to

247 reserve boundaries for OGMAs. Any remaining cedar-hemlock forest patches in OGMAs after

248 this merging of layers that were smaller than 0.1 ha in size were removed from our dataset, again

249 to ensure that we were not inflating estimates of due to differences in

250 spatial resolution (when biogeoclimatic zone and boundaries for OGMAs were overlaid for

251 spatial analysis). Boundaries for OGMAs were converted to lines and buffered by 35, 80 and 120

252 m edge effect thresholds and were used to remove the edges of cedar-hemlock forest stands

253 within OGMAs. Polygon areas and perimeters were calculated in ArcMap using the calculate

254 geometry function. Mean Nearest Neighbour Distance was calculated using the Average Nearest

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255 Neighbor tool. The proportion of OGMAs within 120 m of past cutblocks (harvest years 1980 -

256 2017) was assessed to determine the exposure of current OGMAs to edge effects.

257 Forest management terminology used in the present discussion follows the British

258 Columbia Forest Practices Board (https://www.bcfpb.ca/news-resources/glossary/).

259

260 Results

261 Cedar-hemlock forests (all ages, including past cutblocks) cover 426,925 ha in the

262 Kispiox Timber Harvesting Landbase (excluding lands in provincial parks), with 50% of these

263 currently mapped as age class 8 (141 to 250 years in age) and 9 (> 250 years in age) stands (Fig.

264 1). Within the central-Kispiox region, oldDraft cedar-hemlock forest stands represent 59% of the

265 present-day landscape, down from an estimated pre-harvest value of 87% (Table 1). The mean

266 and largest patch size for old cedar-hemlock forest stands in the central-Kispiox has fallen

267 dramatically from preharvest values, by 91 and 82% respectively (Table 1).

268 In the central-Kispiox, 18% of OGMAs containing cedar-hemlock forest stands are

269 currently adjacent to harvest cutblocks on one side or more (Fig. 2A). An example of this pattern

270 of adjacent cutblocks can be seen for OGMAs in the detailed central-Kispiox map area (Fig. 2B),

271 where edge effects extend inward along most of the eastern boundary of highly elongate

272 OGMAs.

273 The overall proportion of old-forests in the four unlogged Kispiox watersheds examined

274 was 85% (Table 1). The landscape shape index of old-forest patches in the post-harvest central-

275 Kispiox area was more than twice that of the pre-harvest estimate. The mean patch size for old

276 cedar-hemlock forest stands in remaining unlogged watersheds varied considerably from

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277 drainage to drainage, lowest in the narrow and more confined Kuldo Creek valley, highest in the

278 wider Sicintine River valley.

279 The landscape shape index for OGMAs containing old cedar-hemlock forest patches was

280 6.7, indicating that most OGMAs were markedly elongate in shape (Table 2). Similarly, the

281 landscape shape index of the old cedar-hemlock forest patches within OGMAs was 6.4, again

282 indicating highly elongate shapes.

283 Analysis of edge effects confirms this conclusion about patch shapes, with “interior”

284 habitats in OGMAs reduced to 19,838, 11,720, and 7,754 ha respectively, after 35, 80, and 120

285 m edge effect thresholds were applied (Table 2). As the edge effect threshold increases, 286 remaining interior forest habitat patchesDraft become less elongate in shape, with landscape shape 287 index values only 3.0 for the 120 m edge effect. Imposition of a 120 m edge effect creates many

288 smaller and discrete patches of old cedar-hemlock forest in the Kispiox, with a mean patch size

289 of 11 ha. The largest patch index value for old cedar-hemlock forest patches in OGMAs was

290 1.067. The mean nearest neighbour distance was 1.35 for OGMAs containing old cedar-hemlock

291 forest patches, falling to 0.46 km after imposition of 35m edge effects.

292 The majority of old cedar-hemlock forest patches within OGMAs (62%) were smaller

293 than 20 ha in size, although cumulatively, these small patches represented only 9.8% of the total

294 area of old cedar-hemlock forest patches in OGMAs (Fig. 5A). Few old cedar-hemlock forest

295 patches in OGMAs were over 100 ha in size (8.9% of total); however, these patches constituted

296 the majority (58%) of the area protected in OGMAs. With the imposition of edge effects in

297 calculations (Figs. 5B to 5D), the amount of old cedar-hemlock forest in OGMAs over 100 ha in

298 size declined by over 80%. Of the interior old-forest ICH patches retained after imposition of a

299 120 m edge effect, 74% were smaller than 5 ha in size.

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300 Over 95% of old-forest patches in the historical central-Kispiox landscape were 1,000 ha

301 or larger (Fig. 6B), declining to 64% in the present-day landscape (Fig. 6A). Present-day central-

302 Kispiox landscapes have a much higher number of old-forest patches under 5 ha in size (69%),

303 than did historical (preharvest) landscapes (37%). Of the remaining unlogged watersheds, the

304 Sicintine, Upper Skeena, and Shedin each have a high proportion of old-forest area in patches

305 over 1,000 ha in size (84, 98, and 96% respectively) (Figs. 6D, 6E, and 6F respectively). The

306 narrower Kuldo Creek watershed had a lower proportion of 100+ ha old-forest patches, at 50%

307 (Fig. 6C).

308 Detailed examination of the shape and size of individual OGMAs within four

309 representative study areas (see Fig. 4 – areas A to D) confirms the generally elongate shape of

310 old cedar-hemlock forest patches withinDraft individual OGMAs2. The majority of the OGMAs found

311 within the landscape area Fig. 4A (four out of five individual OGMAs examined) contain only

312 “edge” habitats after applying a 120 m edge effect threshold. Twenty-four percent of the area

313 within the largest Old-Growth Management Area found in Fig. 4B remained interior habitat after

314 imposition of a 120 m edge effect threshold. In contrast, OGMAs created as a buffer alongside

315 the Skeena River (Fig. 4C) are highly elongate in shape, with little interior habitat (<25%)

316 remaining after application of a 120 m edge (applied from the land side only). Old-Growth

317 Management Areas placed around small watersheds that included cedar-hemlock forests at their

318 lower margin (Fig. 4D) provided one of the more effective shapes for reducing potential edge

319 effect exposure.

320

2 Detailed maps of the Old-Growth Management Areas outlined in areas A, B, C, and D in Fig. 4 are available in supplemental materials A to E. Landscape metrics for these areas are provided in supplementary materials H. Page 15

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321 Discussion

322

323 Old-forest dependant lichen taxa and Old-Growth Management Areas

324 Given the projected loss of much of the current habitat utilized by old-forest dependant

325 canopy lichen communities in the Kispiox, it is essential that where lands are set aside for

326 conservation they will be suitable for that purpose. This is particularly important in context of

327 planned stand rotation age in the Kispiox (age of forest stands at the next harvest), which

328 commonly occurs between 80 to 120 years after stand establishment in wet cedar-hemlock

329 forests in B.C. (Stevenson et al. 2011). This age threshold will largely preclude old-forest

330 dependant lichens from establishing in second-growthDraft forests, which require stands of 200-250

331 years in age or more to establish and persist (Campbell and Fredeen 2004; Price et al. 2017).

332 Old-Growth Management Area reserves are one of the major land use designations

333 available in B.C. to conserve old-forest habitats. Although logging is not allowed in OGMAs,

334 other resource uses such as aggregate extraction and construction of forest access roads are

335 allowed (Environmental Law Centre 2013). The designation of old-forest reserves within

336 managed forests occurs in many jurisdictions, including old-growth Conservation Reserves in

337 Ontario (Ontario Ministry of Natural Resources and Forestry 2014), Conservation Zones in

338 Quebec (Messier et al. 2009), Protected Forest Areas in Sweden (Hedwall and Mikusiński 2015),

339 Woodland Key Habitats in northern Europe (Aune et al. 2005; Timonen et al. 2011), and to some

340 degree, different types of green-tree retention patches (Gustafsson et al. 2012).

341 As harvesting progresses over the next four to five decades in the Kispiox, old-forest

342 stands will increasingly be limited to areas set aside as spatially defined OGMAs. These reserves

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343 currently represent 7.1% of Kispiox cedar-hemlock forests (30,433 out of 426,925 ha; Table 2

344 and results above). Although old-forests will be retained elsewhere (outside OGMAs under

345 aspatial retention targets), current indications are that OGMAs will provide the majority of future

346 habitat for old-forest dependant oceanic lichens, given their preference for high nutrient status

347 valley-bottom old-forest stands (Goward and Burgess 1996). These productive habitats are rarely

348 conserved by aspatial retention targets, which are usually met from stands that are inoperable or

349 otherwise uneconomic to log (Radies et al 2009) and generally have limited habitat value for

350 oceanic lichens (Goward and Burgess 1996; Goward and Spribille 2005).

351

352 Edge effects and forest fragmentationDraft

353 A key consideration in making decisions about protected area designations in the Kispiox

354 must be that of edge effects (Esseen and Renhorn 1998; Kivistö and Kuusinen 2000; Aune et al.

355 2005). Many of the OGMAs within the Kispiox were established as linear corridors alongside

356 streams and rivers (e.g. as in Fig. 4C). While retention of stream-side forests in OGMAs may

357 provide benefits for aquatic biota (Gomi et al. 2006), for canopy lichens and other old-forest

358 dependant taxa, their narrow width means that most trees will be subject to severe edge effects

359 after adjacent forests are logged. Although the stream-side (natural) edge of OGMAs along rivers

360 should be wind-firm and subject to limited edge effects (Moen and Jonsson 2003), edge effects

361 from their land-based side will be pervasive once adjacent stands are logged. Our projections

362 suggest that only 25% of old ICH forests in Kispiox OGMAs will remain interior habitat after

363 surrounding landscapes are fully converted to second-growth forests.

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364 Johansson et al. (2018), similarly found that retained buffer strips along streams in

365 central-Sweden could not support lichen diversity in the decades after harvesting of adjacent

366 stands. Johansson et al. suggest that retained lichen communities in these buffer strips faced a

367 high probability of extinction. The interaction of size and shape of protected areas with edge

368 effects has been noted in many previous studies. Habitat reserves that have less elongate shapes

369 (shorter perimeters compared to the area of patches) are generally less prone to edge effects

370 compared to elongate patches of the same size (Laurance and Yensen 1991). This is a key factor

371 when considering habitat quality for canopy lichens, where fetch effects, such as the penetration

372 of desiccating winds into old-forest patches are more pervasive in reserves with elongate and/or

373 irregular shapes (Essen and Renhorn 1998; Perhans et al. 2009).

374 Landscape shape index values forDraft old cedar-hemlock habitat patches in Kispiox OGMAs,

375 at 6.4, confirms their highly elongate nature, although, these values were lower than most found

376 by Armenteras et al. (2003) in isolated forest reserves in the eastern Andes, where landscape

377 shape indices reached 19.1. Ranta et al. (1998) found that the size and shape of forest fragments

378 was critical to understanding edge effects in Brazilian rain forest reserves. When they applied a

379 100 m edge effect threshold Ranta et al. found that only 27% of the forest patches remained

380 interior habitat, close to the values we project for Kispiox OGMAs.

381 It should be noted that edge effects can impose both direct and indirect sources of

382 mortality for canopy lichens. Direct effects, such as the bleaching of smoker’s lung lichen thalli

383 observed by Gauslaa et al. (2018), can rapidly restrict lichens to interior habitats after edges are

384 created. Burton (2002) found that edge effects from increased irradiance extended at least 65-70

385 m into the canopy after clearcut edges were created in the Kispiox. Indirect sources of mortality

386 after edges are created, such as greater wind-throw, breakage of limbs, and dieback of trees, will

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387 restrict suitable habitat over a much longer time-period, potentially many decades after edges are

388 created (Esseen 1994). An example of cumulative edge effect impacts on canopy lichens is the

389 study of Cameron et al. (2013), who observed that Erioderma pedicellatum (Hue) P.M. Jørg., a

390 boreal rainforest cyanolichen from eastern Canada, was sensitive to edge effects well beyond 100

391 m buffer zones previously established at clearcut edges.

392 While many of the current Kispiox OGMAs were designated as stream-side buffers, the

393 Botrychium Basin Old-Growth Management Area, originally proposed as the Botrychium Basin

394 Sensitive Area (now identified as Old-Growth Management Area reserve SKE_KIS_611),

395 provides an example of a reserve specifically established to protect rare plant communities

396 (Williston 2002; COSEWIC 2006). This reserve includes one of the few western redcedar-

397 leading old-forest stands in the Kispiox Draft(Williston 2002). Recently observations of clearcut

398 logging blocks extending to the northern reserve boundary (Google Earth Timelapse 2018) and

399 proposed adjacent cutblocks (Williston 2002) raise concerns about the loss of interior-forest

400 habitat in this reserve. This is an issue that warrants immediate management intervention,

401 including possible enlargement/amalgamation of adjacent OGMAs, so as to provide a larger area

402 of contiguous old-forest habitat at this site.

403 The impacts of fragmentation of old-forest stands and isolation of OGMAs (from other

404 old-forest patches) in the Kispiox will vary by lichen species. The mean nearest neighbour

405 distance of 1.35 km for OGMAs containing cedar-hemlock forests should allow for the

406 continued dispersal of some lichen groups, such as early-successional Alectoriod lichens

407 (Goward 2003). This distance, however, is well beyond the dispersal ability for old-forest

408 dependant oceanic lichens (Sillett et al. 2000). Direct measurement of lichen soredial dispersal is

409 difficult, however, several lines of evidence suggest that oceanic lichens are highly dispersal

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410 limited. Jüriado et al. (2011), for instance, found that the vegetative dispersal distance between

411 the host trees of L. pulmonaria was only 15–30 m, while Öckinger et al. (2005) found that

412 dispersal of L. pulmonaria soredia was between 35 and 70 m. These limitations of dispersal

413 highlight the importance of providing long-term continuity of forest cover for old-forest

414 dependant lichens (Fritz et al. 2008; Scheidegger and Werth 2009; Dymytrova et al. 2018).

415 The predicted mean future patch size for old cedar-hemlock forests in Kispiox OGMAs

416 (43 ha before edge effects are considered, 11 ha after considering edge effects) is worrying in

417 this regard. Young and Mitchell (1994), working in wet temperate New Zealand forests,

418 suggested that most old-forest reserves under 10 ha in size will be dominated by edge effect

419 patterns and processes. We would predict that oceanic lichens in small old forest patches in the

420 Kispiox will be highly vulnerable to localDraft extinction events due to fluctuations in the number of

421 lichen thalli (i.e. metapopulation dynamics) following stochastic disturbance events, such as

422 drought (Belinchón et al. 2017). Given their limited dispersal ability, it is likely that oceanic

423 lichens in Kispiox OGMAs, once isolated, will suffer from severe extinction debt effects

424 (Berglund and Jonsson 2005; Kuussaari et al. 2009).

425 The relatively small projected size of future interior habitat in Kispiox OGMAs as

426 landscape-level logging progresses will also pose serious risks to other old-forest dependant taxa,

427 such as the Northern Goshawk, whose populations have declined precipitously in the Kispiox

428 (Doyle 2015). Doyle notes that 83% of known breeding sites now occur in stands that have been

429 reduced below 100 ha in size by logging, well below the minimum size threshold of 176 ha

430 recommended by McClaren et al. (2015). In this regard, conservation of oceanic lichens in the

431 Kispiox would have significant “umbrella” effects (sensu Roberge and Angelstam 2004) for a

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432 range of taxa (Imbeau and Desrochers 2002; Cushman et al. 2011), due to their shared

433 requirement for extensive areas of contiguous old-forest habitat.

434 Although the imposition of edge effects will vary considerably between OGMAs,

435 depending on the timing and pattern of future harvest entries, current management plans would

436 eventually see all Kispiox OGMAs surrounded by younger forests. The pattern and size of

437 adjacent clearcut harvest blocks can have major impacts on the severity of edge effects. The use

438 of dispersed cutblocks (harvest blocks widely spread throughout the landscape; see Delong 2007

439 for further discussion) can result in disproportionate edge effect impacts (Mladenoff et al. 1993,

440 Fenger 1996). This has led for calls to reconsider the size and shape of dispersed cutblocks, so

441 that logging impacts are concentrated in fewer parts of the landscape, while others can retain

442 larger areas of contiguous natural forestDraft (Bergeron et al. 1999; DeLong 2007; Stevenson et al.

443 2011).

444

445 Other designations for old-forest protection

446 The other major land designation that supports conservation of old-forest stands within

447 the Kispiox is that of Provincial Parks. The major park in the Kispiox containing old cedar-

448 hemlock forests is Swan Lake Kispiox River Provincial Park, in the headwaters of the Kispiox

449 River watershed. This park contains the largest contiguous patch of old cedar-hemlock forests in

450 the region, at over 11,000 ha. Although Swan Lake Kispiox River Provincial Park provides

451 valuable habitat for old-forest dependant taxa, as the sole major large contiguous protected area

452 (for old cedar-hemlock forests) in a landscape of over 2.9 million ha (when considering both the

453 Kispiox and adjacent Nass Timber Supply Areas), it is vulnerable to habitat loss from major

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454 natural disturbance events such as fire, a risk likely to be exacerbated by (Hebda

455 1997). This vulnerability of conserving habitat in single large reserves has been highlighted by

456 numerous conservation studies, epitomized by the acronym SLOSS, i.e. “single large versus

457 several smaller reserves” (Simberloff and Abele 1976; Scheia et al. 2013). A mixed strategy,

458 retaining reserves across the Kispiox landscape, with appropriate buffering from edge effects,

459 would be a more prudent conservation strategy. Clearly, Swan Lake Kispiox River Provincial

460 Park must be complemented by additional conservation measures in the adjacent timber

461 harvesting land base, sensu Hansen and Defries (2007).

462 463 Conclusions and Recommendations Draft 464 The combined influence of edge effects on OGMAs, the small patch size of old cedar-

465 hemlock forests in OGMAs, and the low levels of planned future old-forest retention generally in

466 the Kispiox (especially when set against historic levels of old-forest cover) places old-forest

467 dependant taxa at major risk of extirpation as surrounding landscapes are progressively

468 converted to plantation forestry. As presently implemented, Old Growth Management Areas in

469 the Kipsiox must effectively be regarded as “sinking-ships”, unable to retain viable communities

470 of old-forest dependant lichens into the future. We would recommend several modifications to

471 forest management practices that could avert or slow this projected loss of old-forest dependant

472 taxa in the Kispiox.

473 First and foremost, would be the implementation of 120 m buffer zones around existing

474 Kispiox OGMAs. This measure would eliminate much of the potential for future edge effects. It

475 is important to note that buffer zones do not have to be strict no-harvest zones, rather they can be

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476 zones where partial cut harvesting under specific constraints is allowed, thus maintaining canopy

477 structure next to OGMAs. Stevenson and Coxson (2008) demonstrated that leave-tree retention

478 next to forest edges can create “soft-edges” that ameliorate fetch effects on sensitive canopy

479 lichens such as smoker’s lung lichen. Numerous publications have proposed the adoption of

480 partial cut silvicultural systems to maintain ecosystem properties while providing a sustainable

481 timber supply in wet temperate rainforests (Coates and Burton 1997; Beese et al. 2003; Mitchell

482 and Beese 2011).

483 Another approach that bears consideration, at least in the long-term, is the recruitment of

484 additional old-growth reserves from existing second-growth stands. This strategy has been

485 proposed for the Great Bear Rainforest on B.C.’s central coast, where options for retaining

486 legacy forest stands are limited by past loggingDraft practices (Price et al. 2017). In the Kispiox, this

487 is not yet a critical constraint. We would instead prioritize strategies for the Kispiox that

488 conserve existing high conservation value old cedar-hemlock forest stands, a recommendation

489 first made by Goward and Burgess (1996). The importance of conserving existing old-growth

490 forest stands is highlighted by our growing awareness of the importance of forest continuity to

491 maintaining lichen species richness and abundance (McMullin and Wiersma 2019).

492 Perhaps the most important action that could be taken by Kispiox forest managers and

493 delegated decision-makers would be the designation of a much higher proportion of old cedar-

494 hemlock forest stands in the Kispiox into spatially designated protected areas, using OGMAs or

495 other protected area designations, such as conservancies or Indigenous Protected Areas.

496 Currently identified unlogged watersheds, such as Kuldo and Shedin Creeks, and the Sicintine

497 and upper Skeena River areas, offer significant opportunities for designation of new protected

498 areas. With a current (legally mandated) target for old-forest retention in the Kispiox of 31%, the

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499 designation of new spatially defined reserves could take place without having significant impacts

500 on the long-term timber supply. This would be an important step towards conserving old-forest

501 dependant lichens, allowing Old-Growth Management Areas to truly function as “lifeboats”

502 within the Kispiox landscape.

503

504 Acknowledgements

505 We would like to acknowledge funding support provided by the Ministry of Environment and

506 Climate Change Canada and the University of Northern British Columbia. Advice and assistance

507 from Scott Emmons (UNBC) on the GIS analysis is gratefully acknowledged. We thank Mike 508 Jull (Aleza Lake Research Forest) for reviewingDraft an early draft of the manuscript.

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509 References

510 Alaback, P., Nowacki, G., and Saunders, S. 2013. Natural disturbance patterns in temperate

511 rainforests of southeast Alaska and adjacent British Columbia. Chapter 4. In North

512 Pacific temperate rainforests, ecology and conservation. Edited by G.H. Orians and J.W.

513 Schoen. University of Washington Press, Seattle, WA. Pages 73-88.

514 Armenteras, D., Gast, F., and Villareal, H. 2003. Andean forest fragmentation and the

515 representativeness of protected natural areas in the eastern Andes, Colombia. Biol. Cons.

516 113: 245-256. doi:10.1016/S0006-3207(02)00359-2.

517 Aune, K., Jonsson, B.G., and Moen, J. 2005. Isolation and edge effects among woodland key 518 habitats in Sweden: Is forest policyDraft promoting fragmentation? Biol. Cons. 124: 89-95. 519 doi:10.1016/j.biocon.2005.01.015.

520 Beese, W.J., Dunsworth, B.G., Zielke, K., and Bancroft, B. 2003. Maintaining attributes of old-

521 forests in coastal B.C. through variable retention. For. Chron. 79: 570-578.

522 doi:10.5558/tfc79570-3.

523 Belinchón, R., Harrison, P.J., Mair, L., Várkonyi , G., and Snäll, T. 2017. Local epiphyte

524 establishment and future metapopulation dynamics in landscapes with different

525 spatiotemporal properties. Ecology 98: 741-750. doi:10.1002/ecy.1686

526 Bergeron, Y., Harvey, B., Leduc, A., and Gauthier, S. 1999. Forest management guidelines

527 based on natural disturbance dynamics: Stand- and forest-level considerations. For.

528 Chron. 75: 49-54. doi:10.5558/tfc75049-1.

Page 25

https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 26 of 58

529 Berglund, H., and Jonsson, B.G. 2005. Verifying an extinction debt among lichens and fungi in

530 northern Swedish boreal forests. Conserv. Biol. 19: 338-348. doi:10.1111/j.1523-

531 1739.2005.00550.x.

532 British Columbia Conservation Data Centre. 2019. B.C. Species & Ecosystems Explorer.

533 Available from http://a100.gov.bc.ca/pub/eswp/ (Accessed Sept. 30, 2019).

534 British Columbia Forest Practices Code. 2004. Order establishing provincial non-spatial old-

535 growth objectives. Forest planning and practice regulations.

536 https://www2.gov.bc.ca/assets/gov/farming-natural-resources-and-industry/natural-

537 resource-use/land-water-use/crown-land/land-use-plans-and-objectives/policies- 538 guides/old_growth_order_may18th_final.pdfDraft (Accessed Sept. 30, 2019). 539 British Columbia Forest Practices Board. 2012. Conserving old growth forests in BC.

540 Implementation of old-growth retention objectives under FRPA. Special Investigation.

541 FPB/SIR/36. https://www.bcfpb.ca/wp-content/uploads/2016/05/SIR36-OGMAs.pdf

542 (Accessed Sept. 30, 2019).

543 British Columbia Ministry of Environment and Climate Change Strategy. 2018. BC Parks,

544 Ecological Reserves, and Protected Areas https://catalogue.data.gov.bc.ca/dataset/bc-

545 parks-ecological-reserves-and-protected-areas (Accessed Sept. 30, 2019).

546 British Columbia Ministry of Forests and Range. 2008. Kispiox Timber Supply Area rationale

547 for Allowable Annual Cut (AAC) determination. Effective January 1, 2008, Jim

548 Snetsinger, Chief Forester. 63 pages. https://www2.gov.bc.ca/assets/gov/farming-natural-

549 resources-and-industry/forestry/stewardship/forest-analysis-inventory/tsr-annual-

550 allowable-cut/kispiox_tsa_rationale.pdf (Accessed Sept. 30, 2019).

Page 26

https://mc06.manuscriptcentral.com/cjfr-pubs Page 27 of 58 Canadian Journal of Forest Research

551 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural

552 Development. 2017a. Forest Tenures. FADM: Timber Supply Area (TSA).

553 https://catalogue.data.gov.bc.ca/dataset/fadm-timber-supply-area-tsa (Accessed Sept. 30,

554 2019).

555 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural

556 Development. 2017b. GeoBC: Digital Road Atlas (DRA)—Master Partially-Attributed

557 Roads. https://catalogue.data.gov.bc.ca/dataset/digital-road-atlas-dra-master-partially-

558 attributed-roads (Accessed Sept. 30, 2019).

559 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural 560 Development. 2018a. Forest AnalysisDraft and Inventory: VRI - Forest Vegetation Composite 561 Polygons and Layer 1. https://catalogue.data.gov.bc.ca/dataset/vri-forest-vegetation-

562 composite-polygons-and-layer-1 (Accessed Sept. 30, 2019).

563 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural

564 Development. 2018b. Forest Tenures: Forest Tenure Road Section Lines.

565 https://catalogue.data.gov.bc.ca/dataset/forest-tenure-road-section-lines (Accessed Sept.

566 30, 2019).

567 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural

568 Development. 2018c. Forest Analysis and Inventory: Harvested Areas of BC

569 (Consolidated Cutblocks). https://catalogue.data.gov.bc.ca/dataset/harvested-areas-of-bc-

570 consolidated-cutblocks- (Accessed Sept. 30, 2019).

571 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural 572 Development. 2018d. Resource Planning and Assessment: Old Growth Management

Page 27

https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 28 of 58

573 Areas - Legal – Current. https://catalogue.data.gov.bc.ca/dataset/old-growth- 574 management-areas-legal-current (Accessed Sept. 30, 2019).

575 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural

576 Development. 2018e. Forest Analysis and Inventory: BEC Map.

577 https://catalogue.data.gov.bc.ca/dataset/bec-map (Accessed Sept. 30, 2019).

578 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural 579 Development. 2018f. GeoBC: Freshwater Atlas Stream Network. 580 https://catalogue.data.gov.bc.ca/dataset/freshwater-atlas-stream-network (Accessed Sept. 581 30, 2019).

582 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural 583 Development. 2019. Xsu gwin lik"l"inswx:Draft West Babine Sustainable Resource 584 Management Plan. https://www2.gov.bc.ca/gov/content/industry/crown-land-water/land-

585 use-planning/regions/skeena/westbabine-srmp (Accessed Sept. 30, 2019).

586 Burton, P.J. 2002. Effects of clearcut edges on trees in the Sub-Boreal Spruce zone of northwest-

587 central British Columbia. Silva Fenn. 36: 329–352.

588 Cameron, R.P., Neily, T., and Clapp, H. 2013. Forest harvesting impacts on mortality of an

589 endangered lichen at the landscape and stand scales. Can. J. For. Res. 43: 507-511.

590 doi:10.1139/cjfr-2012-0452.

591 Campbell, J., and Fredeen, A. 2004. Lobaria pulmonaria abundance as an indicator of

592 macrolichen diversity in Interior Cedar-Hemlock forests of east-central British

593 Columbia. Can. J. Bot. 82: 970-983. doi:10.1139/b04-074.

594 CANFOR. 2006. Sustainable Forest Management Plan. Tree Farm Licence 30 Canadian Forest

595 Products Ltd. Prince George Operations & BC Timber Sales, Prince George Business

Page 28

https://mc06.manuscriptcentral.com/cjfr-pubs Page 29 of 58 Canadian Journal of Forest Research

596 Area. June 16, 2006.

597 https://www.canfor.com/documents/environmental/plans/TFL30_SFM_Plan_Jun_2006.p

598 df (Accessed Sept. 30, 2019).

599 Cantú-Salazar, L., and Gaston, K.J. 2010. Very Large Protected Areas and Their Contribution to

600 Terrestrial Biological Conservation. BioScience 60: 808–818.

601 doi:10.1525/bio.2010.60.10.7.

602 Coates, D., and Burton, P.J. 1997. A gap-based approach for development of silvicultural

603 systems to address ecosystem management objectives. For. Ecol. Manage.99: 337-354.

604 doi:10.1016/S0378-1127(97)00113-8.

605 Coates, K.D., Banner, A., Steventon, J.D.,Draft LePage, P. and Bartemucci, P. 1997. The Date Creek 606 silvicultural systems study in the Interior Cedar-Hemlock forest of northwestern British

607 Columbia : overview and treatment summaries. Land Management Handbook 38. British

608 Columbia Ministry of Forests. Research Branch. Victoria, B.C.

609 http://bvcentre.ca/files/SORTIE-ND_reports/Coates_et_al_1997_Date_Creek_Lmh38.pdf

610 (Accessed Sept. 30, 2019).

611 Coxson, D., Goward, T., and Werner, J. 2019. The Inland Temperate Rainforest and Interior

612 Wetbelt Biomes of western North America. Earth Systems and Environmental Sciences.

613 Elsevier Scientific. In-press.

614 COSEWIC 2006. COSEWIC assessment and update status report on the cryptic paw Nephroma

615 occultum in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa.

616 https://wildlife-species.canada.ca/species-risk-

Page 29

https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 30 of 58

617 registry/virtual_sara/files/cosewic/sr_cryptic_paw_lichen_e.pdf (Accessed Sept. 30,

618 2019).

619 COSEWIC 2018. COSEWIC Status Report on Smoker’s Lung Lichen Lobaria retigera in

620 Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa.

621 https://wildlife-species.canada.ca/species-risk-

622 registry/virtual_sara/files/cosewic/srSmokersLungLichen2018e.pdf (Accessed Sept. 30,

623 2019).

624 Cushman, S.A., Raphael, M.G., Ruggiero, L.F., Shirk, A.S., Wasserman, T.N., and O’Doherty,

625 E.C. 2011. Limiting factors and landscape connectivity: the American marten in the 626 Rocky Mountains. Landscape EcologyDraft 26: 1137-1149. doi:10.1007/s10980-011-9645-8. 627 DellaSala, D.A., P. Alaback, T. Spribille, H. von Wehrden, and Nauman, R.S. 2011. Just what

628 are temperate and boreal rainforests? Chapter 1. In: Temperate and boreal rainforests of

629 the world: ecology and conservation. Edited by DA DellaSala et al., pages 1-43. Island

630 Press. Washington, D.C..

631 DeLong, S.C. 2007. Implementation of natural disturbance-based management in northern

632 British Columbia. Forest Chron. 83: 326-227. doi:10.5558/tfc2014-011.

633 Doyle, F. 2015. Occupancy and status of Northern Goshawk breeding areas in the Coast

634 Mountains (Kalum), Nadina and Skeena Stikine Resource Districts. Unpublished report.

635 Wildlife Dynamics Consulting. P.O. Box 3596. Smithers, B.C. V0J 2N0. December

636 2015. 22 pages.

637 ftp://142.34.228.79/DND/external/!publish/FSP/2016%20FSP%20Expectation%20Suppo

Page 30

https://mc06.manuscriptcentral.com/cjfr-pubs Page 31 of 58 Canadian Journal of Forest Research

638 rting%20Information/Goshawk%20Breeding%20Area%20Status%202015c%20(Tipping

639 %20Point%20Reached).pdf (Accessed Sept. 30, 2019).

640 Dymytrova L., Brändli, U.B., Ginzler, C., and Scheidegger, C. 2018. Forest history and epiphytic

641 lichens: Testing indicators for assessing forest autochthony in Switzerland. Ecological

642 Indicators 84: 194-207. doi:10.1016/j.ecolind.2017.08.009

643 Environmental Law Centre. 2013. An old-growth protection act for British Columbia. Murray

644 and Anne Fraser Building University of Victoria P.O. Box 1700 STN CSC Victoria, BC,

645 Canada V8W 2Y2.

646 http://www.elc.uvic.ca/press/documents/AnOldGrowthProtectionActforBC- 647 2013Apr10.pdf (Accessed Sept. Draft30, 2019). 648 Esseen, P.A. 1994. Tree mortality patterns after experimental fragmentation of an old-growth

649 conifer forest. Biol. Cons. 68: 19-28. doi:10.1016/0006-3207(94)90542-8.

650 Esseen, P.E., and Coxson, D.S. 2015. Lichens in forest ecosystems. In Handbook of forest

651 ecology. Edited by K. Peh, R. Corlett and Y. Bergeron. Routledge Handbooks. Taylor &

652 Francis Group Ltd. Oxford, U.K..

653 Essen, P.A., and Renhorn, K.E. 1998. Edge effects on an epiphytic lichen in fragmented forests.

654 Conserv. Biol. 12: 1307–1317. doi:10.1111/j.1523-1739.1998.97346.x.

655 Fenger, M. 1996. Implementing biodiversity conservation through the British Columbia Forest

656 Practices Code. For. Ecol. Manage.85: 67-77. doi:10.1016/S0378-1127(96)03751-6.

Page 31

https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 32 of 58

657 Franklin, J.F., Mitchell, R.J., and Palik, B.J. 2007. Natural disturbance and stand development

658 principles for ecological forestry. United States Department of Agriculture Forest

659 Service. Gen. Tech. Rep. NRS-19. Newtown Square, PA..

660 Fritz, O., Gustafsson, L., and Larsson, K. 2008. Does forest continuity matter in conservation? –

661 A study of epiphytic lichens and bryophytes in beech forests of southern Sweden. Biol.

662 Cons. 141: 655-668. doi:10.1016/j.biocon.2007.12.006

663 Gauslaa, Y., and Solhaug, K.A. 1996. Differences in the susceptibility to light stress between

664 epiphytic lichens of ancient and young boreal forest stands. Functional Ecology 10: 344-

665 354. doi:10.2307/2390282.

666 Gauslaa, Y., Lie, M., Solhaug. K.A. andDraft Ohlson, M. 2006. Growth and ecophysiological 667 acclimation of the foliose lichen Lobaria pulmonaria in forests with contrasting light

668 climates. Oecologia 147: 406-416. doi:10.1007/s00442-005-0283-1.

669 Gauslaa, Y, Bartemucci, P., and Solhaug, K.A. 2018. Forest edge-induced damage of cephalo-

670 and cyanolichens in inland old-growth rainforest of northern British Columbia. Can. J.

671 For. Res. 49: 434-439. doi:10.1139/cjfr-2018-0187.

672 Gehlhausen, S.M., Schwartz, M.W., and Augspurger, C.K. 2000. Vegetation and microclimatic

673 edge effects in two mixed-mesophytic forest fragments. Plant Ecology 147: 21–35. doi:

674 10.1023/A:1009846507652.

675 Government of Canada. 2016. Statistics Canada - Census Boundary Files.

676 https://www12.statcan.gc.ca/census-recensement/2011/geo/bound-limit/bound-limit-

677 2016-eng.cfm (Accessed Sept. 30, 2019).

Page 32

https://mc06.manuscriptcentral.com/cjfr-pubs Page 33 of 58 Canadian Journal of Forest Research

678 Gomi, R., Moore, D., and Dhakal, A.S. 2006. Headwater stream temperature response to

679 clear‐cut harvesting with different riparian treatments, coastal British Columbia, Canada.

680 Water Resources Research 42: 1-11. doi:10.1029/2005WR004162.

681 Google Earth Timelapse. 2018. Image center at latitude 55.478170°N, longitude -

682 127.902244°W. https://earthengine.google.com/timelapse/ (Accessed Sept. 30, 2019).

683 Goward, T. 2003. On the Dispersal of Hair Lichen (Bryoria) in High-elevation Oldgrowth

684 Conifer Forests. Canadian Field Naturalist 117: 44-48.

685 Goward, T. 1994. Notes on old-growth-dependant epiphytic macrolichens in inland British

686 Columbia, Canada. Acta Bot. Fenn. 150: 31-38.

687 Goward, T., and Burgess, D. 1996. EpiphyticDraft macrolichens as indicators of forest antiquity in the

688 Kispiox valley (ICHmc zone), with recommendations for the designation of special

689 management areas. Unpublished report prepared for Prince Rupert Forest Region, Bag

690 5000, Smithers, B.C. V0J 2N0.

691 https://www.waysofenlichenment.net/public/pdfs/Goward_Burgess_1996_Oldgrowth_in

692 dicators.pdf (Accessed Sept. 30, 2019).

693 Goward, T., and Spribille,T. 2005. Lichenological evidence for the recognition of inland rain

694 forests in western North America. J. Biogeogr. 32: 1209–1219. doi: 10.1111/j.1365-

695 2699.2005.01282.x.

696 Gustafsson, L., Kouki, J., and Sverdrup-Thygeson, A. 2010. Tree retention as a conservation

697 measure in clearcut forests of northern Europe: a review of ecological consequences.

698 Scandinavian Journal of Forest Research 25: 295-308.

699 doi:10.1080/02827581.2010.497495.

Page 33

https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 34 of 58

700 Gustafsson, L., Baker, S.C., Bauhus, J., Beese, W.J., Brodie, A., Kouki, J., Lindenmayer, D.B.,

701 Lõhmus, A., Pastur, G.M., Messier,C., Neyland, M., Palik, B., Sverdrup-Thygeson, A.,

702 Volney, W.J.A.,Wayne, A., Franklin, J.F. 2012. Retention forestry to maintain

703 multifunctional forests: a world perspective. Bioscience 62: 633-645.

704 doi:10.1525/bio.2012.62.7.6.

705 Hansen, A.J., and Defries, R. 2007. Ecological mechanisms linking protected areas to

706 surrounding lands. Ecological Applications 17: 974-988. doi:10.1890/05-1098.

707 Hansen, A.J., Spies, T.A., Swanson, F.J, and Ohmann, J.L. 1991. Conserving biodiversity in

708 managed forests. BioScience 41: 382-392. doi:10.2307/1311745.

709 Hebda, R.J. 1997. Impact of climate change on biogeoclimatic zones of British Columbia and

710 Yukon. In Responding to globalDraft climate change in British Columbia and Yukon, Vol. 1.

711 Edited by E. Taylor and B Taylor. B.C. Ministry of Environment, Lands and Parks,

712 Victoria, B.C. http://publications.gc.ca/collections/Collection/En56-119-1997E.pdf

713 (Accessed Sept. 30, 2019).

714 Hedwall, P.A., and Mikusiński, G. 2015 Structural changes in protected forests in Sweden:

715 implications for conservation functionality. Can. J. For. Res. 45: 1215–1224.

716 doi:10.1139/cjfr-2014-0470.

717 Hilmo, O., and Holien, H. 2002. Epiphytic lichen response to the edge environment in a boreal

718 Picea abies Forest in central Norway. The Bryologist: 105: 48-56. doi:10.1639/0007-

719 2745(2002)105[0048:ELRTTE]2.0.CO;2.

720 Hofmeister, J., Hosek, J., Malıcek, J., Palice, Z., Syrovatkova, L., Steinova, J., and Cernajova, I.

721 2016. Large beech (Fagus sylvatica) trees as ‘lifeboats’ for lichen diversity in central

Page 34

https://mc06.manuscriptcentral.com/cjfr-pubs Page 35 of 58 Canadian Journal of Forest Research

722 European forests. Biodiversity Conservation 25: 1073-1090. doi:10.1007/s10531-016-

723 1106-x.

724 Imbeau, L., and Desrochers, A. 2002. Area sensitivity and edge avoidance: the case of the Three-

725 Toed Woodpecker (Picoides tridactylus) in a managed forest. Forest Ecology and

726 Management 164: 249-256. doi: 10.1016/S0378-1127(01)00598-9.

727 Johansson, V., Wikström, C.J., and Hylander, K. 2018. Time-lagged lichen extinction in retained

728 buffer strips 16.5 years after clearcutting. Biological Conservation 225: 53-65.

729 doi:10.1016/j.biocon.2018.06.016.

730 Jüriado, I., Liira, J., Csencsics, D., Widmer, I., Adolf, C., Kohv, K., and Scheidegger, C. 2011.

731 Dispersal ecology of the endangered woodland lichen Lobaria pulmonaria in managed

732 hemiboreal forest landscape. BiodiversityDraft and Conservation 8:1803-1819.

733 doi:10.1007/s10531-011-0062-8.

734 Kivistö, L., and Kuusinen, M. 2000. Edge effects on the epiphytic lichen flora of Picea Abies in

735 Middle Boreal Finland. The Lichenologist 32: 387-398. doi:10.1006/lich.2000.0282.

736 Kuussaari, M., Bommarco, R., Heikkinen, R.K., Helm, A., Krauss, J., Lindborg, R., Öckinger,

737 E., Pärtel, Pino, M.J., Rodà, F., Stefanescu, C., Teder, T., Zobel, M., and Steffan-

738 Dewenter, I. 2009. Extinction debt: a challenge for biodiversity conservation. Trends in

739 Ecology & Evolution 24: 564-571. doi:10.1016/j.tree.2009.04.011.

740 Laurance, W.F. and Yensen, E. 1991. Predicting the impacts of edge effects in fragmented

741 habitats. Biol. Cons. 55: 77-92. doi:10.1016/0006-3207(91)90006-U.

742 Lundström, J., Jonsson, F., Perhans, K., and Gustafsson, L. 2013. Lichen species richness on

743 retained aspens increases with time since clearcutting. For. Ecol. Manage. 293: 49-56.

744 doi:10.1016/j.foreco.2012.12.027.

Page 35

https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 36 of 58

745 Marchak, P., Aycock, S., and Herbert, D. 1999. Falldown: forest policy in British Columbia.

746 Ecotrust Canada, Vancouver, B.C. 198 pages.

747 Matveinen-Huju, K., Niemelä, J., Rita, H., and O’Hara, R.B. 2006. Retention-tree groups in

748 clearcuts: Do they constitute ‘life-boats’ for spiders and carabids? For. Ecol.

749 Manage.230: 119-135. doi:10.1016/j.foreco.2006.04.025.

750 McClaren, E., Mahon, T., Doyle, F., Harrower, and William, L. 2015. Science-based guidelines

751 for managing Northern Goshawk breeding areas in coastal British Columbia. Journal of

752 Ecosystems and Management 15: 1-93. https://jem-

753 online.org/index.php/jem/article/viewFile/576/506 (Accessed Sept. 30, 2019).

754 McMullin, R.T., and Wiersma, Y.F. 2019.Draft Out with OLD growth, in with ecological 755 continNEWity: new perspectives on forest conservation. Front. Ecol. Environ.

756 doi:10.1002/fee.2016

757 Meidinger, D. and Pojar, J. 1991. Ecosystems of British Columbia. B.C. Ministry of Forests,

758 Special Report Series 6: 1-330, Victoria.

759 https://www.for.gov.bc.ca/hfd/pubs/Docs/Srs/Srs06.htm (Accessed Sept. 30, 2019).

760 Messier, M., Tittler, R., Kneeshaw, D.D., Gélinas, N., Paquette, A., Berninger, K., Rheault, H.,

761 Meek, P., and Beaulieu, N. 2009. TRIAD zoning in Quebec: Experiences and results after

762 5 years. For. Chron. 85: 885-896. doi:10.5558/tfc85885-6.

763 Mitchell, S.J., and Beese, W.J. 2011. The retention system: reconciling variable retention with

764 the principles of silvicultural systems. For. Chron. 78: 397-403. doi: 10.5558/tfc78397-3.

Page 36

https://mc06.manuscriptcentral.com/cjfr-pubs Page 37 of 58 Canadian Journal of Forest Research

765 Mladenoff, D.J., White, M.A., Pastor, J., and Crow, T.R. 1993. Comparing spatial pattern in

766 unaltered old-growth and disturbed forest landscapes. Ecol. Appl. 3: 294-306.

767 doi:10.2307/1941832.

768 Moen, J., and Jonsson, B.G. 2003. Edge effects on liverworts and lichens in forest patches in a

769 mosaic of boreal forest and . 17: 380-388.

770 doi.org/10.1046/j.1523-1739.2003.00406.x.

771 Mori, A.S., and Kitagawa, R. 2014. Retention forestry as a major paradigm for safeguarding

772 forest biodiversity in productive landscapes: A global meta-analysis. Biol. Cons. 175: 65-

773 73. doi:10.1016/j.biocon.2014.04.016.

774 Öckinger, E., Niklasson, M., and Nilsson,Draft S.G. 2005. Is local distribution of the epiphytic lichen 775 Lobaria pulmonaria limited by dispersal capacity or habitat quality? Biodiversity and

776 Conservation. 14: 759-773. doi: 10.1007/s10531-004-4535-x.

777 Ontario Ministry of Natural Resources and Forestry. 2014. Ontario’s protected areas planning

778 manual, 2014 Edition. Parks and Protected Areas Policy Section, Ministry of Natural

779 Resources, Peterborough, Ont.

780 https://dr6j45jk9xcmk.cloudfront.net/documents/4787/prov-man-planningmanual-app-uc-

781 final-s.pdf (Accessed Sept. 30, 2019).

782 Perhans, K., Appelgren, L., Jonsson, F., Nordin, U., Söderströma, B., and Gustafsson, L. 2009.

783 Retention patches as potential refugia for bryophytes and lichens in managed forest

784 landscapes. Biol. Cons. 142: 1125-1133. https://doi.org/10.1016/j.biocon.2008.12.033.

Page 37

https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 38 of 58

785 Price, K., Lilles, E.B., and Banner, A. 2017 Long-term recovery of epiphytic communities in the

786 Great Bear Rainforest of coastal British Columbia. For. Ecol. Manage.391:296-308.

787 doi:10.1016/j.foreco.2017.02.023.

788 Radies, D.N., Coxson, D.S., Johnson, C.J., and Konwicki, K.. 2009. Predicting canopy

789 macrolichen diversity and abundance within old-growth inland temperate rainforests. For.

790 Ecol. Manage.259:86–97. doi:10.1016/j.foreco.2009.09.046.

791 Ranta, P., Blom, T., Niemela, J., Joensuu, E., and Siitonen, M. 1998. The fragmented Atlantic

792 rain forest of Brazil: size, shape and distribution of forest fragments. Biodiver. Conserv.

793 7: 385-403. doi:10.1023/A:100888581.

794 Roberge, J.M. and Angelstam, P. 2004. DraftUsefulness of the Umbrella Species Concept as a 795 Conservation Tool. Conservation Biology 18: 76-85. doi:10.1111/j.1523-

796 1739.2004.00450.x.

797 Scheia, F.H., Blom, H.H., Gjerd, I., Grytnes, J.A., Heegaard, E., and Sætersdal, M. 2013.

798 Conservation of epiphytes: Single large or several small host trees? Biol. Cons. 168: 144-

799 151. doi:10.1016/j.biocon.2013.10.001

800 Scheidegger, C., and Werth, S. 2009. Conservation strategies for lichens: insights from

801 population biology. Fungal Biology Reviews 23: 55-66. doi:10.1016/j.fbr.2009.10.003

802 Sillett, S.C., McCune, B., Peck, J.E., Rambo, T.R, and Ruchty, A. 2000. Dispersal limitations of

803 epiphytic lichens result in species dependant on old‐growth forests. Ecol. Appl. 10: 789-

804 799. doi:10.1890/1051-0761(2000)010[0789:DLOELR]2.0.CO;2.

Page 38

https://mc06.manuscriptcentral.com/cjfr-pubs Page 39 of 58 Canadian Journal of Forest Research

805 Simberloff, D., and Abele, L.G. 1976. Island biogeography theory and conservation practice.

806 Science 191: 285-286.

807 Snetsinger, J. 2008. Kispiox Timber Supply Area rationale for Allowable Annual Cut (AAC)

808 Determination. British Columbia Ministry of Forests and Range. Victoria, B.C.

809 https://www2.gov.bc.ca/assets/gov/farming-natural-resources-and-

810 industry/forestry/stewardship/forest-analysis-inventory/tsr-annual-allowable-

811 cut/kispiox_tsa_rationale.pdf (Accessed Sept. 30, 2019).

812 Stevenson, S.K., and Coxson, D.S. 2008. Growth responses of Lobaria retigera to forest edge

813 and canopy structure in the inland temperate rainforest, British Columbia. Forest Ecology 814 and Management: 256:618–223.Draft doi:10.1016/j.foreco.2008.05.025. 815 Stevenson, S.K., Armleder, H., Arsenault, A., Coxson, D., DeLong, C., and Jull, M. 2011.

816 Ecology, conservation, and management of British Columbia’s Inland Rainforest. UBC

817 Press. Vancouver, B.C. 432 pages.

818 Suttles, W., and Sturtevant, W.C. 1990. Handbook of North American Indians, Volume 7,

819 Northwest Coast Wayne Suttles, volume editor; William C. Sturtevant, general editor.

820 Smithsonian Institution, Washington, D.C. 1990. xv + 777 pages.

821 Timberline. 2007. Kispiox Timber Supply Area Timber Supply Review III, Timber Supply

822 Analysis Report Version 5.0 Prepared by: Timberline Forest Inventory Consultants Ltd.

823 Prince George, BC. 154 pp. https://www2.gov.bc.ca/assets/gov/farming-natural-

824 resources-and-industry/forestry/stewardship/forest-analysis-inventory/tsr-annual-

825 allowable-cut/kispiox_tsa_analysis_report.pdf (Accessed Sept. 30, 2019).

Page 39

https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 40 of 58

826 Timonen, J., Gustafsson, L., Kotiaho, J.S., and Mönkkönen, M. 2011. Hotspots in cold climate:

827 Conservation value of woodland key habitats in boreal forests. Biol. Cons. 144: 2061-

828 2067. doi:10.1016/j.biocon.2011.02.016.

829 Williston, P. 2002. Botrychium Basin sensitive area plan. Report to the B.C. Ministry of Forests,

830 Kispiox District, Hazelton. 23 pp.

831 Ylisirniöa, A.L. and Hallikainen, V. 2018. Retention patches maintain diversity of epiphytic and

832 epixylic indicator lichens more effectively than solitary trees. Scand. J. Forest Res. 33:

833 320–331. doi:10.1080/02827581.2017.1415370.

834 Young, Y., and Mitchell, N. 1994. Microclimate and vegetation edge effects in a fragmented 835 podocarp-broadleaf forest in NewDraft Zealand. Biological Conservation 67: 63-72. 836 doi:10.1016/0006-3207(94)90010-8.

837

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838 List of Figures:

839 Figure 1. The distribution of Interior Cedar-Hemlock (ICH) forest stands between 141 and 250

840 years in age (ICH 141-250) and greater than 250 years in age (ICH over 250) in the Kispiox

841 Timber Supply Area (Kispiox), showing the location of B.C. Provincial Parks (Provincial Parks),

842 and forest access roads (Roads). The roaded area with first-pass harvesting in the central-Kispiox

843 region, delineated by outline box A, is shown in greater detail in Figure 2. Major unroaded ICH

844 watersheds (B: Kuldo Creek watershed; C: Upper Skeena River; D: Sicintine River; and E:

845 Shedin Creek) are identified in the circled areas. The location of the Kispiox within the province

846 of British Columbia is indicated by the inset map at top right. The Kispiox base map was

847 obtained from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and

848 Rural Development (2017a) and the locationDraft of BC Parks, Ecological Reserves, and Protected

849 Areas from the British Columbia Ministry of Environment and Climate Change Strategy (2018).

850 Forest age class and ecosystem classification data was obtained from the British Columbia

851 Vegetation Resources Inventory database (British Columbia Ministry of Forests, Lands, Natural

852 Resource Operations and Rural Development 2018a) . The location of forest tenure roads was

853 obtained from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and

854 Rural Development (2018b) and from the Digital Road Atlas (British Columbia Ministry of

855 Forests, Lands, Natural Resource Operations and Rural Development 2017b). Data layers were

856 mapped using ArcMap 10.5.

857 Figure 2. A) The location of forest access roads (Roads), past clearcut harvest cutblocks

858 (Cutblocks since 1980), Interior Cedar-Hemlock forest stands between 141 and 250 years in age

859 (ICH 141-250) and greater than 250 years in age (ICH over 250), and Old-Growth Management

860 Area reserves (OGMAs) within the central-Kispiox watershed (see Figure 1 for location). Inset

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861 Plot B) Existing edge effects are shown in detail for the Old-Growth Management Area

862 (Boundary of OGMAs) at the center of (A), delimiting areas that fall within 120 m of adjacent

863 clearcut harvest blocks (Edge Forest), or more than 120 m from adjacent harvest blocks (Interior

864 Forest). Forest age class data and ecosystem classification was obtained from the British

865 Columbia Vegetation Resources Inventory database (British Columbia Ministry of Forests,

866 Lands, Natural Resource Operations and Rural Development 2018a). The location of forest

867 tenure roads was obtained from the British Columbia Ministry of Forests, Lands, Natural

868 Resource Operations and Rural Development (2018b) and from the Digital Road Atlas (British

869 Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development

870 2017b). Forest cutblock locations were obtained from the British Columbia Ministry of Forests,

871 Lands, Natural Resource Operations andDraft Rural Development (2018c). The location of Old

872 Growth Management Areas was obtained from the British Columbia Ministry of Forests, Lands,

873 Natural Resource Operations and Rural Development (2018d). Data layers were mapped using

874 ArcMap 10.5.

875

876 Figure 3. A) Known occurrences of the old-forest dependant temperate rainforest smokers lung

877 lichen (Lobaria retigera) (Smoker’s Lung Lichen Occurrences) that have been found in B.C.’s

878 central-interior mountain ranges in the Wet and Very Wet Interior Cedar-Hemlock forests (ICH

879 Wet and Very Wet), in the Kispiox and upper Nass region in moist ICH forests (ICH Moist), and

880 in the coastal mountain ranges within submaritime Coastal Western Hemlock forests (CWH

881 Submaritime) (from British Columbia Conservation Data Centre 2019). B) Habitat based

882 population models indicate that the largest proportion of L. retigera thalli by far in British

883 Columbia occurs in the Kispiox region (adapted from COSEWIC 2018). Basemap is from the

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884 BC 2016 Census Boundary File (Government of Canada 2016). Data layers include ICH

885 biogeoclimatic ecosystem classification zone mapping from the British Columbia Ministry of

886 Forests, Lands, Natural Resource Operations and Rural Development (2018e); Smoker’s Lung

887 Lichen Occurrences (British Columbia Conservation Data Centre, 2019); and the location of the

888 Kispiox Timber Supply Area (British Columbia Ministry of Forests, Lands, Natural Resource

889 Operations and Rural Development 2017a). Data layers were mapped using ArcMap 10.5.

890 Figure 4. The location of Old-Growth Management Areas in the Kispiox Timber Supply Area

891 (Kispiox), showing areas within reserves that contain Interior Cedar-Hemlock forests (OGMAs

892 ICH); and areas within reserves that are comprised of other biogeoclimatic zone habitats

893 (OGMAs non-ICH). The OGMAs within outline areas (boxes A through D), are shown in greater

894 detail in supplemental materials A to E.Draft The base map of the Kispiox Timber Supply Area is

895 from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural

896 Development (2017a). Mapped locations for Parks, Ecological Reserves, and Protected Areas

897 were from the British Columbia Ministry of Environment and Climate Change Strategy (2018).

898 Interior cedar-hemlock biogeoclimatic ecosystem classification zone mapping was obtained from

899 the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural

900 Development (2018e). Old Growth Management Area data was from the British Columbia

901 Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018d). The

902 location of forest tenure roads was obtained from the British Columbia Ministry of Forests,

903 Lands, Natural Resource Operations and Rural Development (2018b) and from the Digital Road

904 Atlas (British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural

905 Development 2017b). Data for rivers and streams was obtained from British Columbia Ministry

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906 of Forests, Lands, Natural Resource Operations and Rural Development (2018f). Data layers

907 were mapped using ArcMap 10.5.

908

909 Figure 5. The number and area (ha) of Interior Cedar-Hemlock old-forest (> 140 years in age)

910 patches within Old-Growth Management Areas in the Kispiox Timber Supply Area. Patch sizes

911 were calculated using scenarios of A) no edge effects; B) an edge effect of 35 m; C) an edge

912 effect of 80 m; and D) an edge effect of 120 m from reserve edges. Patch size is categorized

913 within defined size class intervals from < 1 ha to 100+ ha. See Fig. 4 for location of Old-Growth

914 Management Areas. Forest age class and ecosystem classification data was obtained from the 915 British Columbia Vegetation ResourcesDraft Inventory database (British Columbia Ministry of 916 Forests, Lands, Natural Resource Operations and Rural Development 2018b, and Old Growth

917 Management Area data from the British Columbia Ministry of Forests, Lands, Natural Resource

918 Operations and Rural Development (2018d). Data analysis was in ArcMap 10.5.

919

920 Figure 6. The number and area (ha) of Interior Cedar-Hemlock old-forest patches (>140 years in

921 age) in: A) “matrix” landscapes of the present day central-Kispiox region (Current – first pass

922 harvest); B) historic landscapes of the central Kispiox region (historical - preharvest); C) the

923 Kuldo Creek watershed (unlogged); D) the Sicintine River watershed (unlogged); E) the upper

924 Skeena River watershed area (unlogged); and F) the Shedin Creek watershed (unlogged). Patch

925 size is categorized within defined size class intervals from < 1 ha to 1000+ ha. See Fig. 1 for

926 location of landscape study areas. Note the different y-axis scaling on the upper two panels (A)

927 and (B). Forest age class and ecosystem classification data was obtained from the British

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928 Columbia Vegetation Resources Inventory database (British Columbia Ministry of Forests,

929 Lands, Natural Resource Operations and Rural Development 2018a), and Old Growth

930 Management Area data from the British Columbia Ministry of Forests, Lands, Natural Resource

931 Operations and Rural Development (2018d). Data analysis was in ArcMap 10.5.

932

Draft

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933 List of Tables

934 Table 1. Fragmentation indices for Interior Cedar-Hemlock old- and very-old (141+ years in age)

935 forest patches in selected landscapes in the Kispiox Timber Supply Area.

936 Table 2. Fragmentation indices for Interior Cedar-Hemlock (ICH) old- and very-old (141+ years

937 in age) forest patches in Old-Growth Management Areas (OGMAs) within the Kispiox Timber

938 Supply Area. “OGMAs containing ICH” are comprised of all OGMAs that contain ICH forests.

939 “ICH only” patches are comprised exclusively of discrete mapped ICH forest patches within

940 Kispiox OGMAs. Landscape metrics indices were additionally calculated for “ICH only” habitat

941 patches in OGMAs after placement of 35, 80, and 120 m buffers from edges. Draft

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Table 1. Fragmentation indices for Interior Cedar-Hemlock old- and very-old (141+ years in age) forest patches in selected landscapes

in the Kispiox Timber Supply Area.

Metric Interior Cedar-Hemlock Old Forest Patches

Study Areas a.

Kuldo Upper Sicintine Shedine Central Central Creek Skeena RiverDraftCreek Kispiox Kispiox (unlogged) (unlogged) (unlogged) (unlogged) (historical - (current –

preharvest) first-pass

harvest)

number of 101 27 11 31 19 138

patches

largest 35 26 40 82 84 15

patch

index

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mean 119 49.5 296.4 383.3 247 1293 patch size

(ha) landscape 2.2 2.3 2.0 0.8 1.3 1.3 shape index mean 0.553 1.196 0.969Draft0.692 2.1 0.7 nearest neighbour distance

(km) total area 4,996 8,002 4,216 9,052 24,576 16,356 in that patch type

(ha)

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largest 2,521 2,632 1,832 7,395 23,357 4,286

single

patch size

in that

patch type

(ha) Draft a) The study areas for unroaded (unlogged) landscapes were the Kuldo (7172 ha), Upper Skeena (10,064 ha), Sicintine (4572 ha),

and Shedin (9,052 ha) watersheds, and the preharvest central-Kispiox watershed (27,820 ha) (see Fig. 1 for locations). The

central-Kispiox watershed area was also examined as a present-day logged landscape.

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Table 2. Fragmentation indices for Interior Cedar-Hemlock (ICH) old- and very-old (141+ years in age) forest patches in Old-Growth

Management Areas (OGMAs) within the Kispiox Timber Supply Area. “OGMAs containing ICH” are comprised of all OGMAs that contain ICH forests. “ICH only” patches are comprised exclusively of discrete mapped ICH forest patches within Kispiox OGMAs.

Landscape metrics indices were additionally calculated for “ICH only” habitat patches in OGMAs after placement of 35, 80, and 120 m buffers from edges.

Metric Interior Cedar-Hemlock HabitatDraft Patches in OGMA Reservesa. OGMAs ICH only ICH only (35 ICH only (80 ICH only

containing (no edge m edge m edge (120 m

ICH (no effect) effect) effect) edge

edge effect) effect) number of 562 712 1736 1354 694 patches largest patch 1.067 0.645 0.565 0.487 0.425 index

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mean patch 92 43 11 9 11

size (ha)

landscape 6.7 6.4 5.6 3.9 3.0

shape index

mean nearest 1.35 1.01 0.46 0.49 0.78

neighbour distance (km) Draft

total area in 51,578 30,433 19,838 11,720 7,754

that patch type

(ha)

largest single 2,280 1,378 1,208 1,040 908

patch size in

that patch type

(ha)

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a) The study areas for OGMA reserves containing ICH Habitat Patches was the area of age class 8 (141-250 yrs in age) and 9

(>250 yrs in age) ICH stands (213,736 ha) contained within the Kispiox TSA (1,301,220 ha).

Draft

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Draft

Figure 1. The distribution of Interior Cedar-Hemlock (ICH) forest stands between 141 and 250 years in age (ICH 141-250) and greater than 250 years in age (ICH over 250) in the Kispiox Timber Supply Area (Kispiox), showing the location of B.C. Provincial Parks (Provincial Parks), and forest access roads (Roads). The roaded area with first-pass harvesting in the central-Kispiox region, delineated by outline box A, is shown in greater detail in Figure 2. Major unroaded ICH watersheds (B: Kuldo Creek watershed; C: Upper Skeena River; D: Sicintine River; and E: Shedin Creek) are identified in the circled areas. The location of the Kispiox within the province of British Columbia is indicated by the inset map at top right. The Kispiox base map was obtained from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2017a) and the location of BC Parks, Ecological Reserves, and Protected Areas from the British Columbia Ministry of Environment and Climate Change Strategy (2018). Forest age class and ecosystem classification data was obtained from the British Columbia Vegetation Resources Inventory database (British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development 2018a) . The location of forest tenure roads was obtained from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018b) and from the Digital Road Atlas (British

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Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development 2017b). Data layers were mapped using ArcMap 10.5.

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Figure 2. A) The location of forest access roads (Roads), past clearcut harvest cutblocks (Cutblocks since 1980), Interior Cedar-Hemlock forest stands between 141 and 250 years in age (ICH 141-250) and greater than 250 years in age (ICH over 250), and Old-Growth Management Area reserves (OGMAs) within the central-Kispiox watershed (see Figure 1 for location). Inset Plot B) Existing edge effects are shown in detail for the Old-Growth Management Area (Boundary of OGMAs) at the center of (A), delimiting areas that fall within 120 m of adjacent clearcut harvest blocks (Edge Forest), or more than 120 m from adjacent harvest blocks (Interior Forest). Forest age class data and ecosystem classification was obtained from the British Columbia Vegetation Resources Inventory database (British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development 2018a). The location of forest tenure roads was obtained from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018b) and from the Digital Road Atlas (British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development 2017b). Forest cutblock locations were obtained from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018c). The location of Old Growth Management Areas was obtained from the British Columbia Ministry of Forests,

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Lands, Natural Resource Operations and Rural Development (2018d). Data layers were mapped using ArcMap 10.5.

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Figure 3. A) Known occurrences of the old-forest dependant temperate rainforest smokers lung lichen (Lobaria retigera) (Smoker’s Lung Lichen Occurrences) that have been found in B.C.’s central-interior mountain ranges in the Wet and Very Wet Interior Cedar-Hemlock forests (ICH Wet and Very Wet), in the Kispiox and upper Nass region in moist ICH forests (ICH Moist), and in the coastal mountain ranges within submaritime Coastal Western Hemlock forests (CWH Submaritime) (from British Columbia Conservation Data Centre 2019). B) Habitat based population models indicate that the largest proportion of L. retigera thalli by far in British Columbia occurs in the Kispiox region (adapted from COSEWIC 2018). Basemap is from the BC 2016 Census Boundary File (Government of Canada 2016). Data layers include ICH biogeoclimatic ecosystem classification zone mapping from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018e); Smoker’s Lung Lichen Occurrences (British Columbia Conservation Data Centre, 2019); and the location of the Kispiox Timber Supply Area (British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development 2017a). Data layers were mapped using ArcMap 10.5.

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Figure 4. The location of Old-Growth Management Areas in the Kispiox Timber Supply Area (Kispiox), showing areas within reserves that contain Interior Cedar-Hemlock forests (OGMAs ICH); and areas within reserves that are comprised of other biogeoclimatic zone habitats (OGMAs non-ICH). The OGMAs within outline areas (boxes A through D), are shown in greater detail in supplemental materials A to E. The base map of the Kispiox Timber Supply Area is from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2017a). Mapped locations for Parks, Ecological Reserves, and Protected Areas were from the British Columbia Ministry of Environment and Climate Change Strategy (2018). Interior cedar-hemlock biogeoclimatic ecosystem classification zone mapping was obtained from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018e). Old Growth Management Area data was from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018d). The location of forest tenure roads was obtained from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018b) and from the Digital Road Atlas (British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development 2017b). Data for rivers and streams was obtained from British Columbia

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Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018f). Data layers were mapped using ArcMap 10.5.

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Figure 5. The number and area (ha) of Interior Cedar-Hemlock old-forest (> 140 years in age) patches within Old-Growth Management Areas in the Kispiox Timber Supply Area. Patch sizes were calculated using scenarios of A) no edge effects; B) an edge effect of 35 m; C) an edge effect of 80 m; and D) an edge effect of 120 m from reserve edges. Patch size is categorized within defined size class intervals from < 1 ha to 100+ ha. See Fig. 4 for location of Old-Growth Management Areas. Forest age class and ecosystem classification data was obtained from the British Columbia Vegetation Resources Inventory database (British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development 2018b, and Old Growth Management Area data from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018d). Data analysis was in ArcMap 10.5.

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Figure 6. The number and area (ha) of Interior Cedar-Hemlock old-forest patches (>140 years in age) in: A) “matrix” landscapes of the present day central-Kispiox region (Current – first pass harvest); B) historic landscapes of the central Kispiox region (historical - preharvest); C) the Kuldo Creek watershed (unlogged); D) the Sicintine River watershed (unlogged); E) the upper Skeena River watershed area (unlogged); and F) the Shedin Creek watershed (unlogged). Patch size is categorized within defined size class intervals from < 1 ha to 1000+ ha. See Fig. 1 for location of landscape study areas. Note the different y-axis scaling on the upper two panels (A) and (B). Forest age class and ecosystem classification data was obtained from the British Columbia Vegetation Resources Inventory database (British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development 2018a), and Old Growth Management Area data from the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (2018d). Data analysis was in ArcMap 10.5.

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