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Mountain pine beetle (MPB, Dendroctonus ponderosae) is a significant mortality agent of Pinus, and climate-driven range expansion is occurring. Pinus defenses in recently invaded areas, including high elevations, are predicted to be lower than in areas with longer term MPB presence. MPB was recently observed in high-elevation forests of the Great Basin (GB) region, North America. Defense and susceptibility in two long-lived species, GB bristlecone pine (Pinus longaeva) and foxtail pine (P. balfouriana), are unclear, although they are sympatric with a common MPB host, limber pine (P. flexilis). We surveyed stands with sympatric GB bristlecone–limber pine and foxtail–limber pine to determine relative MPB attack susceptibility and constitutive defenses. MPB-caused mortality was extensive in limber, low in foxtail and absent in GB bristlecone pine. Defense traits, including constitutive monoterpenes, resin ducts and wood density, were higher in GB bristlecone and foxtail than in limber pine. GB bristlecone and foxtail pines have relatively high levels of constitutive defenses which make them less vulnerable to climate-driven MPB range expansion relative to other highelevation pines. Long-term selective herbivore pressure and exaptation of traits for tree longevity are potential explanations, highlighting the complexity of predicting plant–insect interactions under climate change.

Great Basin bristlecone pine (Pinus longaeva) at 3 sites in western North America near the upper elevation limit of tree growth showed ring growth in the second half of the 20th century that was greater than during any other 50-year period in the last 3,700 years. The accelerated growth is suggestive of an environmental change unprecedented in millennia. The high growth is not overestimated because of standardization techniques, and it is unlikely that it is a result of a change in tree growth form or that it is predominantly caused by CO₂ fertilization. The growth surge has occurred only in a limited elevational band within [almost equal to]150 m of upper treeline, regardless of treeline elevation. Both an independent proxy record of temperature and high-elevation meteorological temperature data are positively and significantly correlated with upper-treeline ring width both before and during the high-growth interval. Increasing temperature at high elevations is likely a prominent factor in the modern unprecedented level of growth for Pinus longaeva at these sites.

This study develops the use of ‘blue rings’ (BR), reflecting incomplete cell wall lignification, as a sensitive thermal indicator in bristlecone pine (Pinus longaeva D.K. Bailey). Using double-stained anatomical thin-sections, we explore the climatic and topographical constraints governing BR formation by developing a time-series from 83 cores and comparing BR occurrence with the full temporal span of available climatic data (1895–2008 CE). Lignification is temperature-dependent and continues at a cellular level post-radial growth completion. As BRs reflect incomplete lignification, they can serve as a higher resolution and more sensitive proxy for past cooling than previously established tree-growth indicators. Results indicate that blue ring formation is primarily induced by low September temperatures and responds more sensitively to cooling than the well-established frost-ring record. Additionally, the occurrence and intensity of blue rings decreases gradually below the upper tree line. Bristlecone pine BRs are demonstrated to have significant capacity to enhance the reconstruction of past cooling events in North America connected with both localized and hemispheric scale forcing over multi-millennial timescales. Given its unmatched longevity, the species offers an unparalleled potential for Holocene length climate reconstruction. Findings also highlight the potential for blue rings to provide a more nuanced understanding of past temperature fluctuations across multi-millennial timescales.

Plants resist herbivores and pathogens by using constitutive (baseline) and inducible (change in defense after an attack) defenses. Inducibility has long been predicted to trade off with constitutive defense, reflecting the economic use of resources. However, empirical evidence for such tradeoffs is variable, and we still lack understanding about when and where defense trade-offs occur. We tested for tradeoffs between constitutive and induced defenses in natural populations of three species of long-lived pines (Pinus balfouriana, P. flexilis, P. longaeva) that differ greatly in constitutive defense and resistance to mountain pine beetle (MPB, Dendroctonus ponderosae). We also assessed how climate influenced constitutive and inducible defenses. At seven high-elevation sites in the western U.S., we simulated MPB attack to induce defenses and measured concentrations of terpene-based phloem defenses on days 0, 15, and 30. Constitutive and induced defenses did not trade off among or within species. Simulated MPB attack induced large increases in defense concentrations in all species independent of constitutive levels. MPB and its symbiotic fungi typically kill trees and thus could be selective forces maintaining strong inducibility within and among species. The contrasting constitutive concentrations in these species could be driven by the adaptation for specializing in harsh, high-elevation environments (e.g., P. balfouriana and P. longaeva) or by competition (e.g., P. flexilis), though these hypotheses have not been empirically examined. Climate influenced defenses, with the greatest concentrations of constitutive and induced defenses occurring at the coldest and driest sites. The interactions between climate and defenses have implications for these species under climate change.

Bristlecone pine (Pinus longaeva) (PILO) trees exhibit exceptional longevity. Their tree‐ring width (TRW) series offer valuable insights into climatic variability. Maximum latewood density (MXD) typically correlates better with temperature variations than TRW, yet PILO MXD records are non‐existent due to methodological challenges related to tree‐ring structure. Here, we used an X‐ray Computed Tomography (X‐ray CT) toolchain on 51 PILO cores from the California White Mountains to build a chronology that correlates significantly (r = 0.66, p < 0.01) with warm‐season (March‐September) temperature over a large spatial extent. This led to the first X‐ray CT‐based temperature reconstruction (1625–2005 CE). Good reconstruction skill (RE = 0.51, CE = 0.32) shows that extending MXD records across the full length of the PILO archive could yield a robust warm‐season temperature proxy for the American Southwest over millennia. This breakthrough opens avenues for measuring MXD in other challenging conifers, increasing our understanding of past climate further, particularly in lower latitudes. Plain Language Summary Ancient Bristlecone pine trees can live for several millennia and hold invaluable climate information. Their annual rings were used to develop millennium‐length records of the Holocene climate. Maximum latewood density (MXD), which is the highest wood density value in the latewood of a tree ring, has been shown to closely follow summer temperature in different conifer species, but not yet in Bristlecone pine. The gnarly and twisted growth of these ancient trees has presented significant hurdles for MXD analysis. Here we apply an X‐ray Computed Tomography toolchain that allows us to 3D scan through the tissue of a tree ring and to map MXD variations. Using this new technique, we were able to reconstruct warm‐season temperature for the American Southwest back to 1625 CE. With these findings, we are confident that a full‐length reconstruction (back to 2575 BCE) can yield the longest annually resolved temperature reconstruction for this continent. Key Points We present the first X‐ray Computed Tomography‐derived maximum latewood density‐based temperature reconstruction using Bristlecone pine tree cores Bristlecone pine maximum latewood density is a reliable proxy for warm‐season temperature over a large part of the American Southwest Our reconstruction (1625–2005 CE) contains low‐frequency variability and can be prolonged over a large part of the Holocene

The instrumental temperature record is of insufficient length to fully express the natural variability of past temperature. High elevation tree-ring widths from Great Basin bristlecone pine ( Pinus longaeva ) are a particularly useful proxy to infer temperatures prior to the instrumental record in that the tree-rings are annually dated and extend for millennia. From ring-width measurements integrated with past treeline elevation data we infer decadal- to millennial-scale temperature variability over the past 4,500 years for the Great Basin, USA. We find that twentieth century treeline advances are greater than in at least 4,000 years. There is also evidence for substantial volcanic forcing of climate in the preindustrial record and considerable covariation between high elevation tree-ring widths and temperature estimates from an atmosphere–ocean general circulation model over much of the last millennium. A long-term temperature decline of ~−1.1 °C since the mid-Holocene underlies substantial volcanic forcing of climate in the preindustrial record.

The tree-killing mountain pine beetle (Dendroctonus ponderosae Hopkins) is an important disturbance agent of western North American forests and recent outbreaks have affected tens of millions of hectares of trees. Most western North American pines (Pinus spp.) are hosts and are successfully attacked by mountain pine beetles whereas a handful of pine species are not suitable hosts and are rarely attacked. How pioneering females locate host trees is not well understood, with prevailing theory involving random landings and/or visual cues. Here we show that female mountain pine beetles orient toward volatile organic compounds (VOCs) from host limber pine (Pinus flexilis James) and away from VOCs of non-host Great Basin bristlecone pine (Pinus longaeva Bailey) in a Y-tube olfactometer. When presented with VOCs of both trees, females overwhelmingly choose limber pine over Great Basin bristlecone pine. Analysis of VOCs collected from co-occurring limber and Great Basin bristlecone pine trees revealed only a few quantitative differences. Noticeable differences included the monoterpenes 3-carene and D-limonene which were produced in greater amounts by host limber pine. We found no evidence that 3-carene is important for beetles when selecting trees, it was not attractive alone and its addition to Great Basin bristlecone pine VOCs did not alter female selection. However, addition of D-limonene to Great Basin bristlecone pine VOCs disrupted the ability of beetles to distinguish between tree species. When presented alone, D-limonene did not affect behavior, suggesting that the response is mediated by multiple compounds. A better understanding of host selection by mountain pine beetles could improve strategies for managing this important forest insect. Moreover, elucidating how Great Basin bristlecone pine escapes attack by mountain pine beetles could provide insight into mechanisms underlying the incredible longevity of this tree species.

Aims To examine the potential mechanistic predictors of germination and first-year survival in two species of Great Basin sub-alpine trees along an elevation gradient on three soil types. Methods Using a network of experimental gardens, we sowed limber pine and Great Basin bristlecone pine along elevational gradients at three sites on three different soil types. We collected germination and first-year survival data of each species while measuring temperature, soil water content, and other environmental variables to examine the potential predictors of first-year survival in these two species. Results Thanks to consecutive anomalously wet and dry years, we found germination and first-year survival to be largely limited by soil type, soil water content, and precipitation timing. Limber pine germination and survival showed weak negative responses while bristlecone pine germination and survival showed stronger negative responses to temperature. Conclusions Young trees are more sensitive to water limitation than to temperature and soil type has a strong moderating effect on water availability. Precipitation timing affected this availability with winter snowpack being less important in establishment than summer monsoonal rain. These results point to the importance of substrate and understanding limitations on all life stages when attempting to predict species range shifts.

The preference-performance hypothesis states that ovipositing phytophagous insects will select host plants that are well-suited for their offspring and avoid host plants that do not support offspring performance (survival, development and fitness). The mountain pine beetle (Dendroctonus ponderosae), a native insect herbivore in western North America, can successfully attack and reproduce in most species of Pinus throughout its native range. However, mountain pine beetles avoid attacking Great Basin bristlecone pine (Pinus longaeva), despite recent climate-driven increases in mountain pine beetle populations at the high elevations where Great Basin bristlecone pine grows. Low preference for a potential host plant species may not persist if the plant supports favorable insect offspring performance, and Great Basin bristlecone pine suitability for mountain pine beetle offspring performance is unclear. We infested cut bolts of Great Basin bristlecone pine and two susceptible host tree species, limber (P. flexilis) and lodgepole (P. contorta) pines with adult mountain pine beetles and compared offspring performance. To investigate the potential for variation in offspring performance among mountain pine beetles from different areas, we tested beetles from geographically-separated populations within and outside the current range of Great Basin bristlecone pine. Although mountain pine beetles constructed galleries and laid viable eggs in all three tree species, extremely few offspring emerged from Great Basin bristlecone pine, regardless of the beetle population. Our observed low offspring performance in Great Basin bristlecone pine corresponds with previously documented low mountain pine beetle attack preference. A low preference-low performance relationship suggests that Great Basin bristlecone pine resistance to mountain pine beetle is likely to be retained through climate-driven high-elevation mountain pine beetle outbreaks.

Dendroclimatic proxies can be generated from the analysis of wood cellular structures, allowing for a more complete understanding of the physiological mechanisms that control the climatic response of tree species. Century-long (1870-2013) time series of anatomical parameters were developed for Great Basin bristlecone pine ( D.K. Bailey) by capturing strongly contrasted microscopic images through a Confocal Laser Scanning Microscope. Environmental information embedded in wood anatomical series was analyzed in comparison with ring-width series using measures of empirical signal strength. Response functions were calculated against monthly climatic variables to evaluate climate sensitivity of cellular features (e.g., lumen area; lumen diameter) for the period 1950-2013. Calibration-verification tests were used to determine the potential to generate long climate reconstructions from these anatomical proxies. A total of eight tree-ring parameters (two ring-width and six chronologies of xylem anatomical parameters) were analyzed. Synchronous variability among samples varied among tree-ring parameters, usually decreasing from ring-width to anatomical features. Cellular parameters linked to plant hydraulic performance (e.g., tracheid lumen area and radial lumen diameter) showed empirical signal strength similar to ring-width series, while noise was predominant in chronologies of lumen tangential width and cell wall thickness. Climatic signals were different between anatomical and ring-width chronologies, revealing a positive and temporally stable correlation of tracheid size (i.e., lumen and cell diameter) with monthly (i.e., March) and seasonal precipitation. In particular, tracheid lumen diameter emerged as a reliable moisture indicator and was then used to reconstruct total March-August precipitation from 1870 to 2013. Wood anatomy holds great potential to refine and expand dendroclimatic records by allowing estimates of plant physiological adaptations to external stressors. Integrating xylem cellular features with ring-width chronologies can widen our understanding of past climatic variability (including annual extreme events) and improve the evaluation of long-term plant response to drought, especially in connection with future warming scenarios.