Darwyn Coxson’s research while affiliated with University of Northern British Columbia and other places

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Publications (7)


Figure 1. Geographical range of southern Deep-Snow Mountain Caribou in British Columbia showing the extent of the interior wetbelt and Caribou (Inland Temperate) Rainforest. Cross-hatched areas show extirpated herds. In colour online.
Figure 3. Lichen biomass by tree height class interval on a per branch basis (g DM branch −1 ; see A & C) and on a per area basis (kg DM ha −1 ; see B & D), with biomass expressed as dry mass (DM). Measurements were taken at Pinkerton Mountain, British Columbia, for each of Alectoria, Bryoria and foliose lichen functional groups in Abies lasiocarpa (A & B) and Picea engelmannii trees (C & D). Each bar represents the mean ± 1 SE for branches within 2 m height class intervals. Adapted from Campbell & Coxson (2001).
Figure 4. Schematic diagram showing five age-class stages in the development of tree-level hyperabundance in Bryoria fremontii and B. pseudofuscescens (= the high-biomass Bryoria spp.) with increasing stand age within the range of Deep-Snow Mountain Caribou. The inner black canopy represents the defoliated zone where hair lichens establish and thrive above the settled depth of the winter snowpack (the A/B trimline), whereas the outer pale sleeve shows the foliated part of the canopy branches, which decreases geometrically with increasing tree age. The Y axis denotes increasing height above the ground and thus increasing ventilation. Zone A = prolonged snow cover precluding hair lichens; Zone B = zone of periodic Bryoria die-back; Zone C = the region of maximum ventilation and Bryoria hyperabundance. See text for further details on Zones A, B and C. After die-off of old trees occurs, a 'black flame' phase can be seen, where Bryoria is abundant throughout the vertical canopy profile. The development of hyperabundance in Alectoria sarmentosa is similar, albeit extending upwards into the upper canopy only in humid forest settings. In colour online.
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The Manna Effect – a review of factors influencing hair lichen abundance for Canada's endangered Deep-Snow Mountain Caribou ( Rangifer arcticus montanus )
  • Article
  • Full-text available

September 2024

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116 Reads

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1 Citation

The Lichenologist

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Darwyn Coxson

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Canada's endangered Deep-Snow Mountain Caribou (DSC) are endemic to mountainous southern inland British Columbia, where they subsist in winter on an almost exclusive diet of epiphytic hair lichens, especially Bryoria fremontii and B. pseudofuscescens (the high-biomass Bryoria spp.) and Alectoria sarmentosa . Importantly, stand-level hair lichen loadings adequate for the dietary needs of DSC rarely occur in forests younger than c . 120–150 years, an unusual form of old-growth dependence hypothetically linked to certain structural features of old forest ecosystems. Not only does this hypothesis accord well with recent insights into hair lichen ecophysiology, it also allows the formulation of a conceptual ‘hyperabundance’ model for the high-biomass Bryoria spp. and lays the foundation for a similar model for A. sarmentosa . In both cases the models point to a massive standing crop of hair lichens in the overstories of old-growth forests; it is this reservoir that, partly by releasing a constant manna-like rain of thallus fragments into the lower canopy, sustains DSC during the winter half year. The outcome is a sustained-yield system resistant to degradation from overbrowsing, yet vulnerable to fragmentation of old-growth forests by industrial forestry, a process of progressive forage reduction that must ultimately place DSC at risk of winter malnutrition. We conclude that stand-level hair lichen hyperabundance is necessarily an attribute of advanced forest age and, at least in the case of Bryoria , cannot be silviculturally induced in stands younger than c . 120–150 years.

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Can partial-cut harvesting be used to extend the availability of terrestrial forage lichens in late-seral pine-lichen woodlands? Evidence from the Lewes Marsh (southern Yukon) silvicultural systems trial

January 2024

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28 Reads

In northern British Columbia and southern Yukon woodland, caribou forage extensively on terrestrial lichens, predominately mat-forming Cladina species in late-successional pine-lichen woodlands. Many of these stands are now reaching a point in their development where lichen abundance declines as feather-moss mats increase. We evaluated the response of forest floor plant communities in pine-lichen woodlands from the southern Yukon Lewes Marsh partial-cutting trial 8 years after harvesting. Photoplot results documented a major decline (>60% ± 5.6% SE) in the mean surface area of existing large clumps of C. mitis in control (unharvested) treatments, whereas the mean surface area of large C. mitis clumps declined by 28% (±15% SE) in the one-third basal-area removal and showed an increase of 13.5% (±25% SE) in the two-thirds basal-area removal. Line intercept transects documented no changes in overall stand-level lichen abundance between pre-harvest (2012) and post-harvest (2021) measurements, while feather-moss mats and dwarf shrubs showed declines and increases, respectively, in partial-cutting harvest plots. Stand thinning may provide a bridging strategy to extend the period of forage lichen availability in late-seral pine-lichen woodlands, an important consideration in landscapes where increasing severity and frequency of fires are changing the seral-state distribution of caribou habitat.


Terrestrial Lichen Caribou Forage Transplant Success: Year 5 and 6 Results

January 2023

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63 Reads

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3 Citations

Restoration Ecology

Sean B. Rapai

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Duncan McColl

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[...]

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Darwyn Coxson

The southern mountain caribou—a sub‐population of caribou found in British Columbia—is listed on Schedule 1 of the Federal Species at Risk Act as Threatened. Woodland caribou are diet specialists, relying on Cladonia sub‐genus Cladina lichen as a primary food source during winter months. Lichens are burned along with trees and other vegetation during stand replacing wildfire events, a natural disturbance in caribou ranges. In an attempt to accelerate the return of post‐fire forests to productive caribou winter terrestrial lichen habitat, this study examined the survival and cover of three species of transplanted lichens in a post‐wildfire environment in north central British Columbia, Canada, both with and without forest litter amendments. Chlorophyll fluorescence was used to evaluate lichen survival by measuring potential photosynthetic activity. The results of this study demonstrate that transplanted fragments and mats of Cladonia sub‐genus Cladina had survived five and six years after being transplanted within a post‐wildfire environment, and had significantly greater percent cover when compared to the controls. The Fv/Fm results indicated that transplanted lichens survived, regardless of species, propagule type or whether amendments were applied. This article is protected by copyright. All rights reserved.


Why is the rainforest lichen Methuselah’s beard (Usnea longissima) so rare in British Columbia’s inland temperate rainforest?

August 2021

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86 Reads

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7 Citations

Coastal (CTR) and inland temperate rainforests (ITR) in western North America share a rich oceanic lichen flora. The distinctive Methuselah’s beard lichen (Usnea longissima Ach.) is an exception to this pattern of shared distributions, with very few ITR locations. Does this absence reflect dispersal limitations or climatic intolerance? To answer this question, we transplanted U. longissima thalli from the CTR to three ITR locations, assessing growth rates against reciprocal CTR transplants. Canopy microclimate measurements provided concurrent data on growth conditions. Growth rate responses (length, mass, and area) were evaluated after summer and full-year transplants. Notwithstanding extended drought conditions during the summer period, annual transplants at two of the three ITR locations supported growth rates comparable to those at the CTR source U. longissima population, with summer dewfall and autumn rains being major ITR hydration sources. Thalli transplanted to a third ITR site (summer measurements only) in a location transitional to the drier interior plateau lost both mass and length. Based on these findings, we suggest that the absence of U. longissima from much of the ITR reflects the combined influence of dispersal limitations and requirements for stands with long site continuity and topographically induced summer wetting of thalli by dewfall.


Figure 3. Potential primary forest (A), current forest (B), and forest loss (C) in the Interior Wetbelt British Columbia (top panel) and Inland Temperate Rainforest (bottom panel), Canada represent percent cover at the watershed scale. Losses were based on the total anthropogenic disturbance layer.
Figure 4. Caribou core habitat, old-growth bird richness concentrations, sensitive fish locations, and potential lichen habitat within the Interior Wetbelt, British Columbia, Canada.
IUCN Red-listed Ecosystem subcriteria/criterion used in the Interior Wetbelt and Inland Temperate Rainforest of British Columbia, Canada. Adapted from Bland et al. [12].
Potential, remaining, and core primary forest in the Interior Wetbelt and Inland Temperate Rainforest, British Columbia, Canada for Red-listed Ecosystem subcriterion A1. Rankings = VU, vulnerable; EN, endangered; CR, critical based on thresholds in Table 1. Adapted from [12].
Combined Red-listed Ecosystem Subcriteria/Criterion A1-C1 and E for the Interior Wetbelt and Inland Temperate Rainforest, BC. Table 1 is split from the Table because the columns were different. NA-not applicable; EN-Endangered, VU-Vulnerable, CR-critical. Adapted from [12].
Red-Listed Ecosystem Status of Interior Wetbelt and Inland Temperate Rainforest of British Columbia, Canada

July 2021

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282 Reads

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10 Citations

The Interior Wetbelt (IWB) of British Columbia, which includes the globally rare Inland Temperate Rainforest (ITR), contains primary forests poorly attributed and neglected in conservation planning. We evaluated the IWB and ITR using four IUCN Red List of Ecosystems Criteria: geographic distribution, environmental degradation (abiotic and biotic factors), and likelihood of ecosystem collapse. Clearcut logging (3.2M ha) represented 57% of all anthropogenic disturbances, reducing potential primary forest by 2.7 million ha (28%) for the IWB and 524,003 ha (39%) for the ITR. Decadal logging rates nearly doubled from 5.3% to 10.2% from 1970s–2000s. Core areas (buffered by 100-m from roads and developments) declined by 70% to 95% for the IWB and ITR, respectively. Vulnerable was assigned to karst, the only abiotic factor assessed, because it was associated with rare plants. For biotic factors, Old-Growth Birds were Vulnerable, Southern Woodland Caribou (Rangifer tarandus caribou) habitat and Sensitive Fish were Endangered, and Old-Growth Lichens habitat was Critical. Overall, the IWB was ranked as Endangered and the ITR as Critical with core area collapse possible within 9 to 18 years for the ITR, considered one of the world’s most imperiled temperate rainforests.


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

April 2020

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114 Reads

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5 Citations

The Kispiox Timber Supply Area, a 1.3 million ha region in northwestern British Columbia, Canada, supports a significant assemblage of temperate rainforest (oceanic) lichens that depend on old forests. Given their known sensitivity to edge effects, we ask whether or not the current configuration of Kispiox Old-Growth Management Area (OGMA) reserves will provide viable future habitat for oceanic lichens as surrounding landscapes are progressively logged in coming decades. Landscape indicators were calculated from provincial map data sets. Old Interior Cedar–Hemlock biogeoclimatic zone forests, the primary habitat for Kispiox oceanic lichens, had a landscape shape index of 6.4 in OGMAs, indicative of elongate shapes susceptible to edge effects. Mean patch size in OGMAs was 43 ha, with the largest patch size being 1 378 ha. In contrast, the landscape shape index for pre-industrial old cedar–hemlock forests was 1.3, with a mean patch size of 1 293 ha and largest patch size of 23 357 ha. When modelled edge effects were extended to 120 m, only 25% of cedar–hemlock forests in Kispiox OGMAs remained interior habitat (7 754 ha total). Adoption of silvicultural practices that maintain buffer zones around existing OGMAs, and the designation of additional OGMAs, especially in watersheds with intact old cedar–hemlock forests, is recommended to conserve oceanic lichen communities in the Kispiox region.


Citations (6)


... Stevenson and Coxson (2007) attributed the increase of Bryoria spp. in partial cuts to increased ventilation rather than increased light. Goward et al. (2024) note the sensitivity of canopy Bryoria communities to reduced ventilation, both within the canopy of individual trees and at a stand level, and the potential for periodic dieback events in wet years within closed-canopy forests. Rambo and North (2012) found no effect of forest thinning on growth of B. fremontii in subalpine forests in California, but this lichen was favoured by lower VPD and greater basal area around trees, buffering canopy microclimate and increasing growth. ...

Reference:

Microclimate drives growth of hair lichens in boreal forest canopies after partial cutting
The Manna Effect – a review of factors influencing hair lichen abundance for Canada's endangered Deep-Snow Mountain Caribou ( Rangifer arcticus montanus )

The Lichenologist

... Experiments involving the transplantation of reindeer lichen on burnt ground have been conducted in boreal regions of northern Sweden (Roturier et al., 2017) and British Columbia, Canada (Rapai et al., 2023), and demonstrated that transplanted fragments survive and establish within a decade of a fire event. The next step forward in ecological restoration is to evaluate whether the system has recovered from disturbance, or is progressing along a trajectory of recovery, including cross-comparison with control and reference ecosystems (Wortley et al., 2013). ...

Terrestrial Lichen Caribou Forage Transplant Success: Year 5 and 6 Results
  • Citing Article
  • January 2023

Restoration Ecology

... At microclimatic scales, growth increases with irradiance during hydration and is often limited by low light in dense forests and lower canopy (Gauslaa et al., 2020;Phinney et al., 2021). Hair lichens have considerable growth potential in optimal climates (McCune et al., 1996;Gauslaa et al., 2007;Phinney et al., 2021;Strother et al., 2022). However, few studies have separated growth from losses by thallus fragmentation and other factors Coxson and Coyle, 2003). ...

Why is the rainforest lichen Methuselah’s beard (Usnea longissima) so rare in British Columbia’s inland temperate rainforest?

... For example, in the province of British Columbia, Canada, which encompasses extensive areas of coastal and interior old-growth forest (DellaSala et al., 2021), temporary deferral of logging has been implemented in a subset of priority remnant old-growth stands. The provincial government implemented these "emergency" deferrals in response to advocacy by communities, Indigenous peoples (termed First Nations in Canada), and non-governmental organizations, and in response to the threat of federal intervention to protect at-risk species [Northern Spotted Owl (Strix occidentalis caurina) and mountain caribou (Rangifer tarandus caribou)] under the Canada's Species At Risk Act (SARA). ...

Red-Listed Ecosystem Status of Interior Wetbelt and Inland Temperate Rainforest of British Columbia, Canada

... Current forestry practices and climate change are predicted to reduce globally significant temperate rainforests in British Columbia and their lichens (Bezzola & Coxson, 2020;Coxson et al., 2019). The Kispiox inland temperate rainforest (hereafter "Kispiox rainforest") supports a rich lichen flora, owing to clean air and an oceanic macroclimate of high humidity and frequent low-intensity summer rain (Coxson et al., 2019). ...

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

... Study site elevation above sea level (a.s.l.) is indicated where data is provided by original authors. The Caribou Rainforest area follows Coxson et al. (2020) While the crucial role played by old-growth conifer forests in providing DSC with winter forage is well established (Environment Canada 2014), the mechanisms involved in the stand-level acquisition of hair lichen hyperabundance remain obscure. At issue is a curious mismatch between, on the one hand, the effective dispersal (Goward 2003b) and early entry into regenerating forests by Alectoria and especially Bryoria (Goward & Campbell 2005;Goward et al. 2022) and, on the other hand, their much-delayed transition into hyperabundance at c. 120-150 years post disturbance Boudreau et al. 2009) or even later (Esseen et al. 1996). ...

The Inland Temperate Rainforest and Interior Wetbelt Biomes of Western North America
  • Citing Chapter
  • January 2019