Special Issue "Ecology and Restoration of Whitebark Pine"

A special issue of Forests (ISSN 1999-4907). This special issue belongs to the section "Forest Ecology and Management".

Deadline for manuscript submissions: closed (30 June 2019).

Special Issue Editors

Dr. Robert E Keane
Website
Guest Editor
Research Ecologist Forest Service, Rocky Mountain Research Station Missoula Fire Sciences Laboratory, Missoula, MT 59808, USA
Interests: wildland fuel science; whitebark pine ecosystem restoration; ecological modeling; climate change science
Dr. Kathryn Ireland
Website
Guest Editor
Assistant Research Professor, Department of Ecology, Montana State University, Bozeman, MT 59717, USA
Dr. Shawn T. McKinney

Guest Editor
Ecologist, University of Montana, Fire Center, and Rocky Mountain Research Station Missoula Fire Sciences Laboratory, Missoula MT 59808
Interests: whitebark pine community ecology; fire ecology; wildlife ecology; climate change

Special Issue Information

Dear Colleagues,

Whitebark pine (Pinus albicaulis) forests have been declining throughout their range in Western North America from the combined effects of mountain pine beetle (Dendroctonus ponderosae) outbreaks, fire exclusion policies, and the exotic disease white pine blister rust. Projected warming and drying trends in climate may exacerbate this decline; however, whitebark pine has a wide climatic tolerance because of its broad distribution coupled with high genetic diversity. The ongoing decline in this high-elevation tree species poses serious consequences for upper subalpine and treeline ecosystems and ecosystem services, and, as a result, whitebark pine is a candidate species for listing under the Endangered Species Act. Many land management agencies are now employing various actions in attempts to restore this valuable forest, including implementing pro-active thinning and burning treatments, planting rust-resistant seedlings, and collecting seed from putative rust resistant trees. However, there has been limited research on this particular species. This Special Issue is designed to be composed of papers that can be used to help restore this important high mountain resource.

Dr. Robert E Keane
Guest Editor

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Keywords

  • Clarks Nutcracker

  • Disturbance

  • Climate Change

  • Mountain Pine Beetle

  • Blister Rust

  • Wildland Fire

  • Forest Management

Published Papers (18 papers)

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Open AccessArticle
Biophysical Gradients and Performance of Whitebark Pine Plantings in the Greater Yellowstone Ecosystem
Forests 2020, 11(1), 119; https://doi.org/10.3390/f11010119 - 19 Jan 2020
Abstract
Research Highlights: The efficacy of planting for restoration is important for ecosystem managers. Planting efforts represent an opportunity for conserving and managing species during a population crisis. Background and Objectives: Federal agencies have been planting whitebark pine (WBP), an important subalpine species that [...] Read more.
Research Highlights: The efficacy of planting for restoration is important for ecosystem managers. Planting efforts represent an opportunity for conserving and managing species during a population crisis. Background and Objectives: Federal agencies have been planting whitebark pine (WBP), an important subalpine species that is late to mature and long-lived, for three decades in the Greater Yellowstone Ecosystem (GYE). These efforts have been met with varying success, and they have not been evaluated beyond the first five years post-planting. Ecosystem managers will continue to plant WBP in the GYE for years to come, and this research helps to inform and identify higher quality habitat during a period of changing climate and high GYE WBP mortality rates. Materials and Methods: We use a combination of field sampling and a water balance model to investigate local biophysical gradients as explanatory variables for WBP performance at twenty-nine GYE planting sites. Results: We found that the WBP growth rate was positively correlated with actual evapotranspiration (AET) and was greatest when cumulative growing season AET was above 350 mm. Growth rate was not strongly affected by competition at the levels found in this study. However, site density change over time was negatively affected by mean growing season temperature and when more than five competitors were present within 3.59 m radius. Conclusions: If they make it to maturity, trees that are planted this season will not begin to produce cones until the latter half of this century. We recommend planting efforts that optimize AET for growth rate objectives, minimize water deficit (WD) that cause stress and mortality, and removing competitors if they exceed five within a short distance of seedlings. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
An Economical Approach to Distinguish Genetically Needles of Limber from Whitebark Pine
Forests 2019, 10(12), 1060; https://doi.org/10.3390/f10121060 - 22 Nov 2019
Abstract
Whitebark pine is difficult to distinguish from limber pine when seed cones are not present. This is often the case because of young stand age, growth at environmental extremes, or harvesting by vertebrate species. Developing an economical genetic identification tool that distinguishes non-cone-bearing [...] Read more.
Whitebark pine is difficult to distinguish from limber pine when seed cones are not present. This is often the case because of young stand age, growth at environmental extremes, or harvesting by vertebrate species. Developing an economical genetic identification tool that distinguishes non-cone-bearing limber from whitebark pine, therefore, could aid many kinds of research on these species. Phylogenetic studies involving limber and whitebark pine suggest that chloroplast DNA sequences differ between these species. We therefore wanted to identify chloroplast loci that could differentiate limber from whitebark pine trees by taking an economical approach involving restriction-site analysis. We generated chloroplast DNA barcode sequences sampled from limber and whitebark pine trees that we identified using attached seed cones. We searched for nucleotide differences associated with restriction endonuclease recognition sites. Our analyses revealed that matK and the psbA-trnH spacer each readily amplified and harbored multiple DNA-sequence differences between limber and whitebark pine. The matK coding sequence of whitebark pine has a BsmAI restriction site not found in limber pine. The psbA-trnH spacer of limber pine has two PsiI restriction sites, neither of which is found in whitebark pine. DNA-sequence and restriction-site analysis of the psbA-trnH spacer from 111 trees showed complete congruence between visually and genetically identified limber (n = 68) and whitebark (n = 43) pine trees. We conclude that restriction site analysis of the chloroplast psbA-trnH spacer and matK involves both minimal technical expertize and research funds. These findings should be of value to foresters interested in species identification and distribution modeling, as well as the analysis of fossil pine pollen, given that gymnosperms transmit chloroplast DNA paternally. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Whitebark Pine in Crater Lake and Lassen Volcanic National Parks: Assessment of Stand Structure and Condition in a Management and Conservation Perspective
Forests 2019, 10(10), 834; https://doi.org/10.3390/f10100834 - 21 Sep 2019
Abstract
Whitebark pine (Pinus albicaulis. Engelm.) is vulnerable to a number of threats including an introduced pathogen (Cronartium ribicola J.C. Fisch.), epidemic levels of native mountain pine beetle (Dendroctonus ponderosae Hopkins), fire suppression, and climate change. To describe the structure of [...] Read more.
Whitebark pine (Pinus albicaulis. Engelm.) is vulnerable to a number of threats including an introduced pathogen (Cronartium ribicola J.C. Fisch.), epidemic levels of native mountain pine beetle (Dendroctonus ponderosae Hopkins), fire suppression, and climate change. To describe the structure of whitebark pine populations in two national parks in the southern Cascades (Crater Lake, Oregon, USA (CRLA) and Lassen Volcanic, California, USA (LAVO) National Parks), we surveyed trees in 30 × 50 × 50 m plots in both parks. We used these plots to describe the extent of white pine blister rust (the disease caused by Cronartium ribicola), mountain pine beetle occurrence, and to elucidate factors influencing the presence of pests and pathogens, cone production, and canopy kill. In each plot, we recorded data related to tree health, including symptoms of blister rust and mountain pine beetle, and reproductive vigor (cone production). In both parks, encroachment from other species, particularly mountain hemlock (Tsuga mertensiana (Bong.) Carrière), was negatively associated with cone production. In CRLA, water stress was a good predictor of blister rust infection and cone production. For CRLA and LAVO, the presence of mountain pine beetle and blister rust was associated with higher canopy kill for whitebark pine. Lastly, we found evidence for a pest-pathogen interaction, mountain pine beetle attack was greater for trees that showed symptoms of blister rust infection in CRLA. Our results indicate that whitebark pine populations in the southern Cascade Range are experiencing moderate levels of blister rust infection compared with other sites across the species range, and that competition from shade-tolerant species may result in an additional threat to whitebark pine in both parks. We present our findings in the context of park management and situate them in range-wide and regional conservation strategies aimed at the protection and restoration of a declining species. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Survival of Whitebark Pine Seedlings Grown from Direct Seeding: Implications for Regeneration and Restoration under Climate Change
Forests 2019, 10(8), 677; https://doi.org/10.3390/f10080677 - 11 Aug 2019
Cited by 2
Abstract
Whitebark pine populations are declining nearly range-wide, primarily from the exotic fungal pathogen that causes white pine blister rust (WPBR). Climate change is expected to exacerbate these declines by decreasing climatically suitable areas. Planting WPBR-resistant seedlings is a key restoration action, but it [...] Read more.
Whitebark pine populations are declining nearly range-wide, primarily from the exotic fungal pathogen that causes white pine blister rust (WPBR). Climate change is expected to exacerbate these declines by decreasing climatically suitable areas. Planting WPBR-resistant seedlings is a key restoration action, but it is costly, time consuming, and labor intensive. Direct seeding—sowing seeds rather than planting seedlings—may reduce costs and open remote areas to restoration; however, its efficacy remains largely unexplored. In this case study, we estimated the annual survival rates (ASR) of seedlings grown from directly sown seeds, and the effect of elevation zone and microsite type on survival. For five years we monitored 184 caches containing one or more seedlings within one study area in the Greater Yellowstone Ecosystem. Seed caches were originally stratified between subalpine forest and treeline and among three microsite types defined by a nurse object: Rocks, trees, and no object. To estimate ASR, we selected the most parsimonious model of a set using AICc. ASR was best described by elevation zone and year and ranged from 0.571 to 0.992. The odds of seedling survival were 2.62 times higher at treeline than in subalpine forest and were 4.6 to 36.2 times higher in 2016–2018 than 2014. We estimated the probability that a whitebark pine seed cache would contain one or more living seedlings six years after sowing to be 0.175 and 0.0584 for treeline and subalpine caches, respectively. We estimated that 1410 and 4229 caches ha−1 would need to be sown at treeline and in subalpine forest, respectively, to attain target restoration densities of 247 established trees ha−1. Our findings, although based on one study area, suggest that climate change may be increasing treeline regeneration, and that direct seeding may be a viable restoration option and climate change mitigation tool for whitebark pine. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Climatic Correlates of White Pine Blister Rust Infection in Whitebark Pine in the Greater Yellowstone Ecosystem
Forests 2019, 10(8), 666; https://doi.org/10.3390/f10080666 - 07 Aug 2019
Abstract
Whitebark pine, a foundation species at tree line in the Western U.S. and Canada, has declined due to native mountain pine beetle epidemics, wildfire, and white pine blister rust. These declines are concerning for the multitude of ecosystem and human benefits provided by [...] Read more.
Whitebark pine, a foundation species at tree line in the Western U.S. and Canada, has declined due to native mountain pine beetle epidemics, wildfire, and white pine blister rust. These declines are concerning for the multitude of ecosystem and human benefits provided by this species. An understanding of the climatic correlates associated with spread is needed to successfully manage impacts from forest pathogens. Since 2000 mountain pine beetles have killed 75% of the mature cone-bearing trees in the Greater Yellowstone Ecosystem, and 40.9% of monitored trees have been infected with white pine blister rust. We identified models of white pine blister rust infection which indicated that an August and September interaction between relative humidity and temperature are better predictors of white pine blister rust infection in whitebark pine than location and site characteristics in the Greater Yellowstone Ecosystem. The climate conditions conducive to white pine blister rust occur throughout the ecosystem, but larger trees in relatively warm and humid conditions were more likely to be infected between 2000 and 2018. We mapped the infection probability over the past two decades to identify coarse-scale patterns of climate conditions associated with white pine blister rust infection in whitebark pine. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Whitebark Pine Recruitment in Sierra Nevada Driven by Range Position and Disturbance History
Forests 2019, 10(5), 455; https://doi.org/10.3390/f10050455 - 25 May 2019
Abstract
Effective restoration of whitebark pine populations will require a solid understanding of factors affecting seedling recruitment success, which may vary by site and biogeographic region. We examined the relationship between whitebark pine seedling recruitment, disturbance history, and range position in three independent studies [...] Read more.
Effective restoration of whitebark pine populations will require a solid understanding of factors affecting seedling recruitment success, which may vary by site and biogeographic region. We examined the relationship between whitebark pine seedling recruitment, disturbance history, and range position in three independent studies in the southern Sierra Nevada, California (CA), USA. In 66 plots broadly distributed across watersheds, we found that whitebark pine seedling density and proportion were greatest at upper elevations, and where canopy cover of whitebark pine was higher (density ranged 0–383 seedlings/ha; x ¯ = 4, σX = 1). Seedling density and proportion were also greater in plots that had recently experienced loss of canopy cover from insects, avalanche, windthrow, or other disturbance effects. In a second study conducted in popular recreational areas, including campgrounds and trailheads, the response of whitebark pine recruitment to disturbance was strongly dependent on the relative position of stands within the range, or proximity to other forest types. Both studies indicated that low to moderate levels of disturbance enhanced whitebark pine recruitment, especially at its range edge, a finding consistent with the early seral status of whitebark observed in previous studies conducted elsewhere in North America. In our third study, a case study at the June Mt. Ski Area, we demonstrate the potential for a downward shift in the whitebark-lodgepole pine ecotone as a result of insect-caused disturbance. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Seed Availability Does Not Ensure Regeneration in Northern Ecosystems of the Endangered Limber Pine
Forests 2019, 10(2), 146; https://doi.org/10.3390/f10020146 - 10 Feb 2019
Cited by 1
Abstract
Research Highlights: When biotic interactions such as disease alter both the seed production capacity of stands, and seedling survivorship, the relative importance of seed availability versus substrate specificity may alter future regeneration opportunities for plant populations. Background and Objectives: We investigated the [...] Read more.
Research Highlights: When biotic interactions such as disease alter both the seed production capacity of stands, and seedling survivorship, the relative importance of seed availability versus substrate specificity may alter future regeneration opportunities for plant populations. Background and Objectives: We investigated the importance of disease severity, seed availability, and substrate limitation to the regeneration dynamics of the endangered limber pine, Pinus flexilis, in two ecosystems with varying forest composition, and different histories of white pine blister rust infection (WPBR; Cronartium ribicola). Materials and Methods: A total of 17 stands from the montane ecoregion (Alberta, Canada) were sampled for seed production between 2007–2010, seedling density, and age structure. Model selection using an information theoretic approach compared a series of a priori models and their combinations, based on our hypotheses on the role biotic variables play in the regeneration process. Results: Despite higher rates of WPBR infection, 2.3 times more seed was available for avian dispersers in the southern ecosystem. Recent seedling regeneration did not correspond to seed production (83 versus 251 seedlings/ha, in southern versus northern ecosystems, respectively), resulting in a seven-fold difference in seed to seedling ratios between ecosystems. Models suggest that disease and vegetation cover were important factors explaining the absence of regeneration in 79.4% of the plots sampled, while basal area (BA) of live limber pine, rocky substrates, ecosystem, South aspects, and slope enhanced limber pine regeneration. Seedling age structures suggest that recent regeneration is less likely in more diseased landscapes, than it was historically (40% versus 72.8% of seedlings < 20 years old, respectively, in southern versus northern ecosystems). Conclusions: At the northern limits of limber pine’s range, seed availability does not ensure regeneration, suggesting that other environmental or biotic factors hinder regeneration. Regeneration was consistently predicted to be lower in the southern ecosystem than in the northern ecosystem, suggesting that natural regeneration and the potential for population recovery are ecosystem dependent. We recommend that monitoring recent seedlings will aid the identification of biotic and abiotic factors affecting regeneration. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Whitebark and Foxtail Pine in Yosemite, Sequoia, and Kings Canyon National Parks: Initial Assessment of Stand Structure and Condition
Forests 2019, 10(1), 35; https://doi.org/10.3390/f10010035 - 07 Jan 2019
Cited by 4
Abstract
The Inventory & Monitoring Division of the U.S. National Park Service conducts long-term monitoring to provide park managers information on the status and trends in biological and environmental attributes including white pines. White pines are foundational species in many subalpine ecosystems and are [...] Read more.
The Inventory & Monitoring Division of the U.S. National Park Service conducts long-term monitoring to provide park managers information on the status and trends in biological and environmental attributes including white pines. White pines are foundational species in many subalpine ecosystems and are currently experiencing population declines. Here we present results on the status of whitebark and foxtail pine in the southern Sierra Nevada of California, an area understudied relative to other parts of their ranges. We selected random plot locations in Yosemite, Sequoia, and Kings Canyon national parks using an equal probability spatially-balanced approach. Tree- and plot-level data were collected on forest structure, composition, demography, cone production, crown mortality, and incidence of white pine blister rust and mountain pine beetle. We measured 7899 whitebark pine, 1112 foxtail pine, and 6085 other trees from 2012–2017. All factors for both species were spatially highly variable. Whitebark pine occurred in nearly-pure krummholz stands at or near treeline and as a minor component of mixed species forests. Ovulate cones were observed on 25% of whitebark pine and 69% of foxtail pine. Whitebark pine seedlings were recorded in 58% of plots, and foxtail pine seedlings in only 21% of plots. Crown mortality (8% in whitebark, 6% in foxtail) was low and significantly higher in 2017 compared to previous years. Less than 1% of whitebark and zero foxtail pine were infected with white pine blister rust and <1% of whitebark and foxtail pine displayed symptoms of mountain pine beetle attack. High elevation white pines in the southern Sierra Nevada are healthy compared to other portions of their range where population declines are significant and well documented. However, increasing white pine blister rust and mountain pine beetle occurrence, coupled with climate change projections, portend future declines for these species, underscoring the need for broad-scale collaborative monitoring. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Landscape Topoedaphic Features Create Refugia from Drought and Insect Disturbance in a Lodgepole and Whitebark Pine Forest
Forests 2018, 9(11), 715; https://doi.org/10.3390/f9110715 - 18 Nov 2018
Cited by 3
Abstract
Droughts and insect outbreaks are primary disturbance processes linking climate change to tree mortality in western North America. Refugia from these disturbances—locations where impacts are less severe relative to the surrounding landscape—may be priorities for conservation, restoration, and monitoring. In this study, hypotheses [...] Read more.
Droughts and insect outbreaks are primary disturbance processes linking climate change to tree mortality in western North America. Refugia from these disturbances—locations where impacts are less severe relative to the surrounding landscape—may be priorities for conservation, restoration, and monitoring. In this study, hypotheses concerning physical and biological processes supporting refugia were investigated by modelling the landscape controls on disturbance refugia that were identified using remotely sensed vegetation indicators. Refugia were identified at 30-m resolution using anomalies of Landsat-derived Normalized Difference Moisture Index in lodgepole and whitebark pine forests in southern Oregon, USA, in 2001 (a single-year drought with no insect outbreak) and 2009 (during a multi-year drought and severe outbreak of mountain pine beetle). Landscape controls on refugia (topographic, soil, and forest characteristics) were modeled using boosted regression trees. Landscape characteristics better explained and predicted refugia locations in 2009, when forest impacts were greater, than in 2001. Refugia in lodgepole and whitebark pine forests were generally associated with topographically shaded slopes, convergent environments such as valleys, areas of relatively low soil bulk density, and in thinner forest stands. In whitebark pine forest, refugia were associated with riparian areas along headwater streams. Spatial patterns in evapotranspiration, snowmelt dynamics, soil water storage, and drought-tolerance and insect-resistance abilities may help create refugia from drought and mountain pine beetle. Identification of the landscape characteristics supporting refugia can help forest managers target conservation resources in an era of climate-change exacerbation of droughts and insect outbreaks. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Whitebark Pine Prevalence and Ecological Function in Treeline Communities of the Greater Yellowstone Ecosystem, U.S.A.: Potential Disruption by White Pine Blister Rust
Forests 2018, 9(10), 635; https://doi.org/10.3390/f9100635 - 12 Oct 2018
Cited by 4
Abstract
In the northern Rocky Mountains of the U.S. and Canada, whitebark pine (Pinus albicaulis Engelm.) is a functionally important species in treeline communities. The introduced fungal pathogen Cronartium ribicola, which causes white pine blister rust, has led to extensive whitebark pine [...] Read more.
In the northern Rocky Mountains of the U.S. and Canada, whitebark pine (Pinus albicaulis Engelm.) is a functionally important species in treeline communities. The introduced fungal pathogen Cronartium ribicola, which causes white pine blister rust, has led to extensive whitebark pine mortality nearly rangewide. We examined four treeline communities within the Greater Yellowstone Ecosystem (GYE) to assess structure and composition, whitebark pine prevalence and functional role, differences in growing season mesoclimate among study areas, and blister rust infection incidence. We found that (1) whitebark pine frequently serves as the majority overall, solitary, and leeward tree island conifer; (2) the prevalence of different tree species in the windward position in tree islands, and thus their potential as tree island initiators, may be predicted from their relative abundance as solitary trees; and (3) white pine blister rust infection incidence ranged from 0.6% to 18.0% across study areas. White pine blister rust poses a threat to treeline development and structure and the provision of ecosystem services in the GYE. Increasing blister rust resistance in nearby subalpine whitebark pine communities through seedling planting or direct seeding projects should eventually result in higher levels of blister rust resistance in whitebark pine in treeline communities. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Effects of Disturbance on Tree Community Dynamics in Whitebark Pine (Pinus albicaulis Engelm.) Ecosystems
Forests 2018, 9(9), 566; https://doi.org/10.3390/f9090566 - 13 Sep 2018
Cited by 3
Abstract
Whitebark pine (Pinus albicaulis Engelm.), an ecologically important tree species in high-elevation ecosystems of western North America, is threatened by white pine blister rust (Cronartium ribicola Fischer) and increased pressure from mountain pine beetle (Dendroctonus ponderosae Hopkins) due to climate [...] Read more.
Whitebark pine (Pinus albicaulis Engelm.), an ecologically important tree species in high-elevation ecosystems of western North America, is threatened by white pine blister rust (Cronartium ribicola Fischer) and increased pressure from mountain pine beetle (Dendroctonus ponderosae Hopkins) due to climate warming. In addition, there is concern that fire suppression may be leading to successional replacement of whitebark by late-seral trees. Despite widespread knowledge that the tree is in decline, there is limited understanding of its successional dynamics, particularly in forests disturbed by white pine blister rust and mountain pine beetle. Our objective was to examine how disturbances have affected forest composition, structure, and seedling regeneration over a 22-year period (1990–2012) at 19 sites in the Cascade Mountains of Washington State (USA). Over that time, 13 sites (68%) were infected by white pine blister rust, 11 (58%) were disturbed by mountain pine beetle, and 5 (26%) experienced wildfire. Tree community composition changed significantly during the study period, primarily due to significant mortality of mature (≥20-cm diameter at breast height) whitebark pine. Despite loss of mature whitebark trees, we found little evidence of successional replacement by other tree species. Whitebark seedling density was unrelated to basal area of mature whitebark pine, but positively correlated with the presence of herb and shrub cover. Our results demonstrate the value of long-term repeated measurements for elucidating successional dynamics. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Designer Niches Promote Seedling Survival in Forest Restoration: A 7-Year Study of Whitebark Pine (Pinus albicaulis) Seedlings in Waterton Lakes National Park
Forests 2018, 9(8), 477; https://doi.org/10.3390/f9080477 - 05 Aug 2018
Cited by 3
Abstract
Designer niches in which environmental variables are controlled are useful in forest restoration to enhance survival of planted tree seedlings. Here, we evaluate particular manipulated habitats, on site variables, and pre-seedling conditions hypothesized to improve the survival rate of whitebark pine (Pinus [...] Read more.
Designer niches in which environmental variables are controlled are useful in forest restoration to enhance survival of planted tree seedlings. Here, we evaluate particular manipulated habitats, on site variables, and pre-seedling conditions hypothesized to improve the survival rate of whitebark pine (Pinus albicaulis) seedlings out-planted in Waterton Lakes National Park. The tree species is in peril due to blister rust and mountain pine beetle infestations in its range; and is a restoration priority in Waterton Lakes because populations in the park are highly infected with blister rust (up to 90%). At Summit Lake, 21 plots were set up and half of each was terra-torched; 1000 seedlings were planted in clusters of three, under four conditions: on burned areas in burned beargrass mats, in burned areas where beargrass mats were not present, in unburned areas where beargrass was present, and in unburned areas without beargrass. This study reports data for the seventh year after planting, and overall, survival was 53% for individual seedlings and at least one seedling survived in 60.8% of clusters. Planting in burned areas increased cluster survival (by 34.3%, p ˂ 0.0001) and planting near microsites increased cluster survival (by 19.3%, p ˂ 0.0001); the type of microsite did not make a difference. Planting in beargrass mats decreased survival, but not significantly (8.9%, p = 0.11) and this was true for burns, not unburned areas. Inoculation with native ectomycorrhizal fungi did not enhance survival most likely because controls on lightly terra-torched and unburned areas had access to local native fungi. This is the first study to report statistics on the planting of seedlings in clusters; the results need to be compared with studies where seedlings are planted individually. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Growth Response of Whitebark Pine (Pinus albicaulis Engelm) Regeneration to Thinning and Prescribed Burn Treatments
Forests 2018, 9(6), 311; https://doi.org/10.3390/f9060311 - 01 Jun 2018
Cited by 3
Abstract
Whitebark pine (Pinus albicaulis Engelm.) forests play a prominent role throughout high-elevation ecosystems in the northern Rocky Mountains, however, they are vanishing from the high mountain landscape due to three factors: exotic white pine blister rust (Cronartium ribicola Fischer) invasions, mountain [...] Read more.
Whitebark pine (Pinus albicaulis Engelm.) forests play a prominent role throughout high-elevation ecosystems in the northern Rocky Mountains, however, they are vanishing from the high mountain landscape due to three factors: exotic white pine blister rust (Cronartium ribicola Fischer) invasions, mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks, and successional replacement by more shade-tolerant tree species historically controlled by wildfire. Land managers are attempting to restore whitebark pine communities using prescribed fire and silvicultural cuttings, but they are unsure if these techniques are effective. The objective of this study was to determine how whitebark pine regeneration responds to selective thinning and prescribed burn treatments. We studied changes in diameter growth after restoration treatments using ring width measurements obtained from 93 trees at four sites in Montana and Idaho that were treated in the late 1990s. Overall, the average annual radial growth rates of the trees in treated areas were greater than those of trees in control areas. Specifically, there were significant increases in the growth ratio (180%) in the two sites that were both thinned and later burned. Younger regeneration showed more response to the treatments than older regeneration. All sites showed high variability in post-treatment growth rates across individual trees, with greater variability for trees in treated areas than in trees from the control areas. Results suggest that whitebark pine regeneration can respond to thin and burn release treatments and that managers may see positive results in areas that are treated similarly. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessFeature PaperArticle
Whitebark Pine in the Northern Cascades: Tracking the Effects of Blister Rust on Population Health in North Cascades National Park Service Complex and Mount Rainier National Park
Forests 2018, 9(5), 244; https://doi.org/10.3390/f9050244 - 03 May 2018
Abstract
Whitebark pine (Pinus albicaulis Engelm.) is a key component of subalpine and alpine ecosystems in the northern Cascades. The species’ survival is threatened by white pine blister rust, mountain pine beetle, fire exclusion, and climate change. We monitored whitebark pine in permanent [...] Read more.
Whitebark pine (Pinus albicaulis Engelm.) is a key component of subalpine and alpine ecosystems in the northern Cascades. The species’ survival is threatened by white pine blister rust, mountain pine beetle, fire exclusion, and climate change. We monitored whitebark pine in permanent plots in two national parks three times between 2004 and 2016. The proportion of live trees showing evidence of blister rust infection increased in North Cascades National Park Service Complex from 32% in 2004 to 51% in 2016 and from 18% to 38% in Mount Rainier National Park. Mortality increased from 7% to 21% in North Cascades National Park Service Complex and 38% to 44% in Mount Rainier National Park. The percent of live infected and dead whitebark pine increased with south and east aspects and mortality decreased with elevation. Annualized mortality rates calculated for the entire study period were 1.5% in Mount Rainier National Park and 2.3% in North Cascades National Park Service Complex. Although these rates decreased between the first time period (2004–2009) and the second time period (2009–2016), the prevalence of infected and dead whitebark pine increased across all park landscapes over time and increased in smaller diameter whitebark pine trees. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Ten Years of Monitoring Illustrates a Cascade of Effects of White Pine Blister Rust and Focuses Whitebark Pine Restoration in the Canadian Rocky and Columbia Mountains
Forests 2018, 9(3), 138; https://doi.org/10.3390/f9030138 - 14 Mar 2018
Cited by 7
Abstract
Whitebark pine forests are declining due to infection by white pine blister rust and mountain pine beetle, combined with the effects of climate change and fire suppression. The Canadian Rocky and Columbia Mountains represent a large portion of the whitebark range; a vast [...] Read more.
Whitebark pine forests are declining due to infection by white pine blister rust and mountain pine beetle, combined with the effects of climate change and fire suppression. The Canadian Rocky and Columbia Mountains represent a large portion of the whitebark range; a vast area, exemplifying the need for knowledge about whitebark pine stands to target restoration. The aim of our work was to identify variables predicting live tree infection, seedling infection, canopy kill, mortality, and regeneration across this region, and present the results in spatially-explicit formats to assist land managers with restoration. Live tree and seedling infection by white pine blister rust increased over the last decade and cascading effects of the disease are intensifying, including canopy kill and mortality. We show that large diameter trees are more likely to be infected, and the highest infection rates are in southern and western areas. The conditions for seedling infection are more strongly influenced by fine-scale climatic conditions than for trees. Areas with low regeneration are: (1) the dry east slopes where live tree infection is low; and (2) where live tree infection rates are high, suggesting that canopy kill and mortality are influencing regeneration. Results highlight where to target restoration and coordinate across boundaries. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessFeature PaperArticle
Pinus albicaulis Engelm. (Whitebark Pine) in Mixed-Species Stands throughout Its US Range: Broad-Scale Indicators of Extent and Recent Decline
Forests 2018, 9(3), 131; https://doi.org/10.3390/f9030131 - 09 Mar 2018
Cited by 10
Abstract
We used data collected from >1400 plots by a national forest inventory to quantify population-level indicators for a tree species of concern. Whitebark pine (Pinus albicaulis) has recently experienced high mortality throughout its US range, where we assessed the area of [...] Read more.
We used data collected from >1400 plots by a national forest inventory to quantify population-level indicators for a tree species of concern. Whitebark pine (Pinus albicaulis) has recently experienced high mortality throughout its US range, where we assessed the area of land with whitebark pine present, size-class distribution of individual whitebark pine, growth rates, and mortality rates, all with respect to dominant forest type. As of 2016, 51% of all standing whitebark pine trees in the US were dead. Dead whitebark pines outnumbered live ones—and whitebark pine mortality outpaced growth—in all size classes ≥22.8 cm diameter at breast height (DBH), across all forest types. Although whitebark pine occurred across 4.1 million ha in the US, the vast majority of this area (85%) and of the total number of whitebark pine seedlings (72%) fell within forest types other than the whitebark pine type. Standardized growth of whitebark pines was most strongly correlated with the relative basal area of whitebark pine trees (rho = 0.67; p < 0.01), while both standardized growth and mortality were moderately correlated with relative whitebark pine stem density (rho = 0.39 and 0.40; p = 0.031 and p < 0.01, respectively). Neither growth nor mortality were well correlated with total stand basal area, total stem density, or stand mean diameter. The abundance, extent, and relative growth vs. mortality rates of whitebark pine in multiple forest types presents opportunities for management to encourage whitebark pine recruitment in mixed-species stands. The lodgepole pine forest type contained more whitebark pine seedlings (35%) than any other forest type, suggesting that this forest type represents a potential management target for silvicultural treatments that seek to facilitate the recruitment of whitebark pine seedlings into larger size classes. National forest inventories in other countries may use a similar approach to assess species of concern. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Open AccessArticle
Historic Frequency and Severity of Fire in Whitebark Pine Forests of the Cascade Mountain Range, USA
Forests 2018, 9(2), 78; https://doi.org/10.3390/f9020078 - 07 Feb 2018
Cited by 1
Abstract
Whitebark pine (Pinus albicaulis Engelm.) is a foundation species of high elevation forest ecosystems in the Cascade Mountain Range of Oregon, Washington, and British Columbia. We examined fire evidence on 55 fire history sites located in the Cascade Range. To estimate dates [...] Read more.
Whitebark pine (Pinus albicaulis Engelm.) is a foundation species of high elevation forest ecosystems in the Cascade Mountain Range of Oregon, Washington, and British Columbia. We examined fire evidence on 55 fire history sites located in the Cascade Range. To estimate dates of historic fires we analyzed 57 partial cross-sections from fire-scarred trees plus 700 increment cores. The resulting 101 fire events indicate fire has been a widespread component of Cascadian whitebark pine stands. Results are site specific and vary considerably. Whitebark pine stands appear to burn in a variety of severities and frequencies. Sites where fire intervals were detected ranged from 9 to 314 years, with a median of 49 years, and averaging 67 years. Fire intervals shortened significantly with higher latitudes. In assessing the most recent fire event at each site, overall, 56 percent burned as stand replacing events. In the 20th century, the number of fires diminished significantly. Due to conservation imperatives, re-introducing fire should be undertaken with extreme care to avoid substantial mortality of this endangered species. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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Review

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Open AccessReview
Managing Wildfire for Whitebark Pine Ecosystem Restoration in western North America
Forests 2018, 9(10), 648; https://doi.org/10.3390/f9100648 - 18 Oct 2018
Cited by 2
Abstract
Wildfire in declining whitebark pine forests can be a tool for ecosystem restoration or an ecologically harmful event. This document presents a set of possible wildfire management practices for facilitating the restoration of whitebark pine across its range in Western North America. These [...] Read more.
Wildfire in declining whitebark pine forests can be a tool for ecosystem restoration or an ecologically harmful event. This document presents a set of possible wildfire management practices for facilitating the restoration of whitebark pine across its range in Western North America. These management actions are designed to enhance whitebark pine resilience and health, while also being effective wildfire management measures. The actions are presented by the three phases of the wildfire continuum: Before, during, and after a wildfire. Current pre-wildfire restoration actions, such as mechanical thinning’s, prescribed burning, and fuel treatments, can also be designed to be fuel treatment activities that allow more effective suppression of wildfires when needed. Three wildfire strategies can be implemented while the wildfire is burning—full suppression, partial suppression, and wildland fire use (letting some fires burn under acceptable conditions)—for protecting valuable whitebark pine trees and for ecosystem restoration. Finally, post-wildfire activities include planting rust-resistant seedlings and monitoring effects of the wildfires. Recommended wildfire management practices for the wildfire continuum are specified by region, site type, and stand type in the last section of this paper. Full article
(This article belongs to the Special Issue Ecology and Restoration of Whitebark Pine)
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