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Bristlecone pine dwarf mistletoe (Arceuthobium microcarpum [Engelm.] Hawksw. & Wiens subsp. aristatae J.M.Scott & Mathiasen) severely parasitizes Rocky Mountain bristlecone pine (Pinus aristata Engelm.) on the San Francisco Peaks, Arizona. We estimated the current incidence of infection and extent of mortality in a mistletoe-infested stand of bristlecone pine on Schultz Peak, a southeast subsidiary ridge of the San Francisco Peaks. Dendrochronological techniques were used to compare radial growth between lightly, moderately, and severely infected and noninfected bristlecone pines. We also paired limber pine (Pinus flexilis James) with noninfected and infected bristlecone pines as another method of estimating the effects of the mistletoe on radial growth. Results suggested that the incidence of infection has increased by approximately 10%, and mortality has increased by approximately 20% since the mid-1970s. Severely infected bristlecone pines had significant growth losses and less annual variation in growth than limber pine or lightly infected and noninfected bristlecone pines. Spatial and temporal trends in mortality of bristlecone pines were measured in two large mortality locations on Schultz Peak. Cross-dating cores from dead bristlecone pines indicated that mortality had rapidly increased starting in 1996. Years of high mortality were related to both high mountain pine beetle (Dendroctonus ponderosae Hopkins) activity and drought conditions.

Like many other high elevation alpine tree species, Rocky Mountain bristlecone pine (Pinus aristata Engelm.) may be particularly vulnerable to climate change. To evaluate its potential vulnerability to shifts in climate, we defined the suitable climate space for each of four genetic lineages of bristlecone pine and for other subalpine tree species in close proximity to bristlecone pine forests. Measuring changes in the suitable climate space for lineage groups is an important step beyond models that assume species are genetically homogenous. The suitable climate space for bristlecone pine in the year 2090 is projected to decline by 74% and the proportional distribution of suitable climate space for genetic lineages shifts toward those associated with warmer and wetter conditions. The 2090 climate space for bristlecone pine exhibits a bimodal distribution along an elevation gradient, presumably due to the persistence of the climate space in the Southern Rocky Mountains and exclusion at mid-elevations by conditions that favor the climate space of other species. These shifts have implications for changes in fire regimes, vulnerability to pest and pathogens, and altered carbon dynamics across the southern Rockies, which may reduce the likelihood of bristlecone pine trees achieving exceptional longevity in the future. The persistence and expansion of climate space for southern bristlecone pine genetic lineage groups in 2090 suggests that these sources may be the least vulnerable in the future. While these lineages may be more likely to persist and therefore present opportunities for proactive management (e.g., assisted migration) to maintain subalpine forest ecosystem services in a warmer world, our findings also imply heighted conservation concern for vulnerable northern lineages facing range contractions.

Using a combination of 23 radiocarbon ages and annual ring counts from 18 Rocky Mountain bristlecone pine (Pinus aristata) remnants above the local present-day limits, a period of higher treeline has been determined for two sites near the Continental Divide in central Colorado. The highest remnants were found about 30 m above live bristlecone pines of similar size. The majority of the remnants, consisting of standing snags, large logs, and smaller remains, are highly eroded, such that the innermost annual rings of all but one are missing. The radiocarbon ages obtained from the oldest wood recovered from each remnant indicate that the majority were established above the present-day limit of bristlecone pine from prior to 2700 cal. yr BP to no later than about 1200 cal. yr BP. These radiocarbon ages combined with the annual ring count from the corresponding remnant indicate that the majority of the sampled remnants grew above the present-day limit of bristlecone pine from sometime before 2700 cal. yr BP to about 800 cal. yr BP. Evidence of recent climatic warming is demonstrated at one of the sites by young bristlecone pine saplings growing next to the highest remnants; the saplings were established after AD 1965 and represent the highest advance of treeline in at least 1200 years.

Natural forests are increasingly invaded by nonnative pests and pathogens that threaten host species with population extirpation and cascading ecological impacts. The regeneration for resilience (R4R) framework provides a decision structure to prioritize limited resources and utilize artificial and natural regeneration management to offer the best likelihood of success in positioning stands and landscapes to support multi-generational self-sustaining host populations in the presence of the nonnative invader by (1) increasing host population size to offset invader-caused mortality, (2) increasing the frequency of genetic resistance traits in host populations in habitats that enable their expression and durability to retard future mortality and facilitate population sustainability and recovery, and (3) maintain host genetic diversity, adaptive capacity, and population connectivity. This application is designed for conditions where the nonnative invader is expected to be a persistent threat, the host species naturally has some genetic resistance to the invasive species, and the forest host populations are integral to support valued ecosystem processes and services. The R4R framework has been developed for application in high elevation five-needle pine ecosystems of North America impacted and threatened by the nonnative pathogen Cronartium ribicola that causes the lethal disease white pine blister rust (WPBR). Several examples using the R4R framework to integrate information on current forest condition, WPBR risk or hazard, genetic resistance to WPBR, and host population dynamics and silvics to prioritize areas and design interventions are discussed. Effective management of forest regeneration dynamics can increase forest resilience and adaptive capacity to mitigate impacts of invasive species.

Trade‐offs between plant defense investment and fitness traits, including growth, are often invoked to explain evolutionary strategies targeted at resisting herbivores. Many Pinus species have specialized herbivores, including the mountain pine beetle (MPB), Dendroctonus ponderosae, and have historically been a focus of defense investigations. We compared defense traits of two high‐elevation Pinus species, P. aristata and P. flexilis, that are hosts to MPB and hypothesized to have different growth and defense traits and potential trade‐offs. Interspecific differences were assessed by sampling trees within the same stands, and intraspecific differences were assessed by sampling stands at sites across latitudes where both species co‐occurred. Constitutive defenses were measured at Day 0, and the timing, concentration, and composition of an induced resin defense response were assessed by sampling at 1, 4, and 30 days following either mechanical wounding only or a simulated MPB attack using its primary fungal symbiont Grosmannia clavigera. At Day 4, induced resin concentrations did not differ between mechanical wounding and simulated MPB attack in either species. By Day 30, resin defense concentrations in response to simulated MPB attack were greater than those in response to mechanical wounding and were >19‐fold greater than constitutive levels. Results suggest that initial induced resin defense responses in the two species are likely generalized, with a delayed response that is targeted specifically at MPB and G. clavigera. At all sites, P. aristata had higher concentrations of constitutive and Day‐30 induced resin defenses than P. flexilis, although P. flexilis induced proportionately more. Trade‐offs in growth and defense between the species were only found at the two most climatically favorable sites where P. aristata grew slower than P. flexilis. No trade‐offs were found between the two defense types at either biological scale. Overall, our findings highlight that the two pine species growing in the same stands (1) have a delayed response to a specialized native herbivore and fungal symbiont, (2) only exhibited interspecific defense–growth trade‐offs at two climatically favorable sites, and showed no intraspecific defense–growth trade‐offs, (3) showed no trade‐offs between constitutive and induced defenses at either biological scale, and (4) have evolved different defense strategies.

Great Basin bristlecone pine (Pinus longaeva) and foxtail pine (Pinus balfouriana) are valuable paleoclimate resources due to their longevity and climatic sensitivity of their annually-resolved rings. Treeline research has shown that growing season temperatures limit tree growth at and just below the upper treeline. In the Great Basin, the presence of precisely dated remnant wood above modern treeline shows that the treeline ecotone shifts at centennial timescales tracking long-term changes in climate; in some areas during the Holocene climatic optimum treeline was 100 meters higher than at present. Regional treeline position models built exclusively from climate data may identify characteristics specific to Great Basin treelines and inform future physiological studies, providing a measure of climate sensitivity specific to bristlecone and foxtail pine treelines. This study implements a topoclimatic analysis-using topographic variables to explain patterns in surface temperatures across diverse mountainous terrain-to model the treeline position of three semi-arid bristlecone and/or foxtail pine treelines in the Great Basin as a function of growing season length and mean temperature calculated from in situ measurements. Results indicate: (1) the treeline sites used in this study are similar to other treelines globally, and require a growing season length of between 147-153 days and average temperature ranging from 5.5°C-7.2°C, (2) site-specific treeline position models may be improved through topoclimatic analysis and (3) treeline position in the Great Basin is likely out of equilibrium with the current climate, indicating a possible future upslope shift in treeline position.

Mountain pine beetle (MPB) (Dendroctonus ponderosae Hopkins) is a native bark beetle that reproduces in pine (Pinus) species across western North America. High population levels can result in widespread host tree mortality. Over the past 2 decades, MPB has been responsible for pine mortality across millions of forested hectares in the western United States. Although a majority of the pine species found in the western United States are considered hosts to MPB, the host status of Rocky Mountain (RM) bristlecone pine (Pinus aristata) is unclear. We surveyed stands across the range of RM bristlecone pine in Colorado, USA, and quantified MPB-caused mortality within the past 10 years in stands where RM bristlecone and at least one other pine species co-occurred. We also evaluated in the field whether successful MPB brood production occurred in RM bristlecone pine. Our results confirm that RM bristlecone pine is susceptible to MPB attack and suitable for MPB reproduction. In mixed-species stands, pine species availability influenced MPB attack occurrence. The proportion of trees experiencing fatal beetle attack within a particular Pinus species was best predicted by the basal area proportion of that species in the stand prior to the most recent 10 years of mortality. These results indicate that RM bristlecone pine is vulnerable to ongoing climate change–induced contact with MPB.

High-resolution pollen, plant macrofossil and magnetic susceptibility (MS) data are presented from an alpine lake sediment core from west-central Colorado, recording changes in vegetation and sedimentation for the latest Pleistocene and Holocene (c. the last 12.5 ka; 1 ka = 1000 cal. yr BP). During the Younger Dryas chron (c. 12.9–11.5 ka), Artemisia steppe or tundra grew around the lake, but by the earliest Holocene (10.7–9.5 ka) a subalpine Picea and Abies parkland was established there. Picea remained important through the early Holocene, but also bristlecone and lodgepole pines (Pinus aristata and P. contorta) grew around the lake. Warming conditions are indicated from 9.5 ka, lasting until c. 4.5–3.5 ka, which may have been the warmest period, with greatest development of monsoonal conditions. Trees subsequently retreated downslope from Kite Lake c. 150–200 m during the last 3.5 ka, establishing their present treeline position. A decrease in total Pinus and increases in Artemisia and piñon (P. edulis) indicate a trend toward progressive climate cooling and enhanced winter precipitation. These long-term climatic trends correlate with Holocene changes in summer insolation.

Aim To understand how tree growth response to regional drought and temperature varies between tree species, elevations and forest types in a mountain landscape. Location Twenty-one sites on an elevation gradient of 1500 m on the San Francisco Peaks, northern Arizona, USA. Methods Tree-ring data for the years 1950-2000 for eight tree species (Abies lasiocarpa var. arizonica (Merriam) Lemm., Picea engelmannii Parry ex Engelm., Pinus aristata Engelm., Pinus edulis Engelm., Pinus flexilis James, Pinus ponderosa Dougl. ex Laws., Pseudotsuga menziesii var. glauca (Beissn.) Franco and Quercus gambelii Nutt.) were used to compare sensitivity of radial growth to regional drought and temperature among co-occurring species at the same site, and between sites that differed in elevation and species composition. Results For Picea engelmannii, Pinus flexilis, Pinus ponderosa and Pseudotsuga menziesii, trees in drier, low-elevation stands generally had greater sensitivity of radial growth to regional drought than trees of the same species in wetter, high-elevation stands. Species low in their elevational range had greater drought sensitivity than co-occurring species high in their elevational range at the pinyon-juniper/ponderosa pine forest ecotone, ponderosa pine/mixed conifer forest ecotone and high-elevation invaded meadows, but not at the mixed conifer/subalpine forest ecotone. Sensitivity of radial growth to regional drought was greater at drier, low-elevation compared with wetter, high-elevation forests. Yearly growth was positively correlated with measures of regional water availability at all sites, except high-elevation invaded meadows where growth was weakly correlated with all climatic factors. Yearly growth in high-elevation forests up to 3300 m a.s.l. was more strongly correlated with water availability than temperature. Main conclusions Severe regional drought reduced growth of all dominant tree species over a gradient of precipitation and temperature represented by a 1500-m change in elevation, but response to drought varied between species and stands. Growth was reduced the most in drier, low-elevation forests and in species growing low in their elevational range in ecotones, and the least for trees that had recently invaded high-elevation meadows. Constraints on tree growth from drought and high temperature are important for high-elevation subalpine forests located near the southern-most range of the dominant species.

Only recently have efforts begun to address how management might prepare currently healthy forests to affect the outcome of invasion by established non-native pests. Cronartium ribicola, the fungus that causes the disease white pine blister rust (WPBR), is among the introductions into North America where containment and eradication have failed; the disease continues to spread. Ecosystem function is impaired by high rust-caused mortality in mature five-needle white pine forests. This paper evaluates five proactive management options to mitigate the development of impacts caused by white pine blister rust in threatened remote high-elevation five-needle pine ecosystems of western North America. They are: reducing pest populations; managing forest composition; improving host vigor; introducing resistant stock with artificial regeneration; and diversifying age class structure to affect the natural selection process for resistance. Proactive intervention to manage and facilitate evolutionary change in the host species may sustain host populations and ecosystem function during pathogen naturalization.