Changes in tree drought sensitivity provided early warning signals to the California drought and forest mortality event.

Climate warming in recent decades has negatively impacted forest health in the western United States. Here, we report on potential early warning signals (EWS) for drought-related mortality derived from measurements of tree-ring growth (ring width index; RWI) and carbon isotope discrimination (∆13 C), primarily focused on ponderosa pine (Pinus ponderosa). Sampling was conducted in the southern Sierra Nevada Mountains, near the epicenter of drought severity and mortality associated with the 2012-2015 California drought and concurrent outbreak of western pine beetle (Dendroctonus brevicomis). At this site, we found that widespread mortality was presaged by five decades of increasing sensitivity (i.e., increased explained variation) of both tree growth and ∆13 C to Palmer Drought Severity Index (PDSI). We hypothesized that increasing sensitivity of tree growth and ∆13 C to hydroclimate constitute EWS that indicate an increased likelihood of widespread forest mortality caused by direct and indirect effects of drought. We then tested these EWS in additional ponderosa pine-dominated forests that experienced varying mortality rates associated with the same California drought event. In general, drier sites showed increasing sensitivity of RWI to PDSI over the last century, as well as higher mortality following the California drought event compared to wetter sites. Two sites displayed evidence that thinning or fire events that reduced stand basal area effectively reversed the trend of increasing hydroclimate sensitivity. These comparisons indicate that reducing competition for soil water and/or decreasing bark beetle host tree density via forest management-particularly in drier regions-may buffer these forests against drought stress and associated mortality risk. EWS such as these could provide land managers more time to mitigate the extent or severity of forest mortality in advance of droughts. Substantial efforts at deploying additional dendrochronological research in concert with remote sensing and forest modeling will aid in forecasting of forest responses to continued climate warming.

Constitutive and Induced Defenses in Long-lived Pines Do Not Trade Off but Are Influenced by Climate.

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.

Oviposition Model for a Southern Population of Mountain Pine Beetle.

We develop a predictive oviposition model for a southern population of mountain pine beetle (MPB) using a previously developed rate curve, incorporating variation in both oviposition rate and fecundity. We also introduce a method for determining the time delay before oviposition. The model describes the probability of oviposition for a season of MPB attacks using hourly phloem temperature and adult MPB attack data. We also develop an asymptotic approximation of MPB oviposition that is much less computationally taxing. The detailed oviposition model and its asymptotic approximation are compared with other ovipositional models for MPB; the predictive capacity of each model is evaluated using previously published observations.

Stronger influence of growth rate than severity of drought stress on mortality of large ponderosa pines during the 2012-2015 California drought.

Forests in the western United States are being subject to more frequent and severe drought events as the climate warms. The 2012-2015 California drought is a recent example, whereby drought stress was exacerbated by a landscape-scale outbreak of western pine beetle (Dendroctonus brevicomis) and resulted in widespread mortality of dominant canopy species including ponderosa pine (Pinus ponderosa). In this study, we compared pairs of large surviving and beetle-killed ponderosa pines following the California drought in the southern Sierra Nevadas to evaluate physiological characteristics related to survival. Inter-annual growth rates and tree-ring stable isotopes (∆13C and δ18O) were utilized to compare severity of drought stress and climate sensitivity in ponderosa pines that survived and those that were killed by western pine beetle. Compared to beetle-killed trees, surviving trees had higher growth rates and grew in plots with lower ponderosa pine basal area. However, there were no detectable differences in tree-ring ∆13C, δ18O, or stable isotope sensitivity to drought-related meteorological variables. These results indicate that differences in severity of drought stress had little influence on local, inter-tree differences in growth rate and survival of large ponderosa pines during this drought event. Many previous studies have shown that large trees are more likely to be attacked and killed by bark beetles compared to small trees. Our results further suggest that among large ponderosa pines, those that were more resistant to drought stress and bark beetle attacks were in the upper echelon of growth rates among trees within a stand and across the landscape.

Management of Western North American Bark Beetles with Semiochemicals.

We summarize the status of semiochemical-based management of the major bark beetle species in western North America. The conifer forests of this region have a long history of profound impacts by phloem-feeding bark beetles, and species such as the mountain pine beetle (Dendroctonus ponderosae) and the spruce beetle (D. rufipennis) have recently undergone epic outbreaks linked to changing climate. At the same time, great strides are being made in the application of semiochemicals to the integrated pest management of bark beetles. In this review, we synthesize and interpret these recent advances in applied chemical ecology of bark beetles for scientists and land managers.

Differential dispersal and the Allee effect create power-law behaviour: Distribution of spot infestations during mountain pine beetle outbreaks.

Mountain pine beetles (MPB, Dendroctonus ponderosae Hopkins) are aggressive insects attacking Pinus host trees. Pines use defensive resin to overwhelm attackers, creating an Allee effect requiring beetles to attack en masse to successfully reproduce. MPB kill hosts, leaving observable, dying trees with red needles. Landscape patterns of infestation depend on MPB dispersal, which decreases with host density. Away from contiguously impacted patches (low beetle densities), infestations are characterized by apparently random spots (of 1-10 trees). It remains unclear whether the new spots are spatially random eruptions of a locally endemic population or a mode of MPB spread, with spatial distribution determined by beetle motility and the need to overcome the Allee effect. To discriminate between the hypothesis of population spread versus independent eruption, a model of spot formation by dispersing beetles facing a local Allee effect is derived. The model gives rise to an inverse power distribution of travel times from existing outbreaks. Using landscape-level host density maps in three study areas, an independently calibrated model of landscape resistance depending on host density, and aerial detection surveys, we calculated yearly maps of travel time to previous beetle impact. Isolated beetle spots were sorted by travel time and compared with predictions. Random eruption of locally endemic populations was tested using artificially seeded spots. We also evaluated the relationship between number of new spots and length of the perimeter of previously infested areas. Spot distributions conformed strongly to predicted power-law behaviour. The spatially random eruption hypothesis was found to be highly improbable. Spot numbers grew consistently with perimeter of previously infested area, suggesting that MPB spread long distances from infestation boundaries via spots following an inverse power distribution. The Allee effect in MPB therefore accelerates, rather than limits, invasion rates, contributing to recent widespread landscape-scale mortality in western North America.

Defense traits in the long-lived Great Basin bristlecone pine and resistance to the native herbivore mountain pine beetle.

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 high-elevation 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.

Reproductive isolation and environmental adaptation shape the phylogeography of mountain pine beetle (Dendroctonus ponderosae).

Chromosomal rearrangement can be an important mechanism driving population differentiation and incipient speciation. In the mountain pine beetle (MPB, Dendroctonus ponderosae), deletions on the Y chromosome that are polymorphic among populations are associated with reproductive incompatibility. Here, we used RAD sequencing across the entire MPB range in western North America to reveal the extent of the phylogeographic differences between Y haplotypes compared to autosomal and X-linked loci. Clustering and geneflow analyses revealed three distinct Y haplogroups geographically positioned within and on either side of the Great Basin Desert. Despite close geographic proximity between populations on the boundaries of each Y haplogroup, there was extremely low Y haplogroup mixing among populations, and gene flow on the autosomes was reduced across Y haplogroup boundaries. These results are consistent with a previous study suggesting that independent degradation of a recently evolved neo-Y chromosome in previously isolated populations causes male sterility or inviability among Y haplotype lineages. Phylogeographic results supported historic contraction of MPB into three separate Pleistocene glacial refugia followed by postglacial range expansion and secondary contact. Distinct sets of SNPs were statistically associated with environmental data among the most genetically distinct sets of geographic populations. This finding suggests that the process of adaptation to local climatic conditions is influenced by population genetic structure, with evidence for largely independent evolution in the most genetically isolated Y haplogroup.

A dynamical model for bark beetle outbreaks.

Tree-killing bark beetles are major disturbance agents affecting coniferous forest ecosystems. The role of environmental conditions on driving beetle outbreaks is becoming increasingly important as global climatic change alters environmental factors, such as drought stress, that, in turn, govern tree resistance. Furthermore, dynamics between beetles and trees are highly nonlinear, due to complex aggregation behaviors exhibited by beetles attacking trees. Models have a role to play in helping unravel the effects of variable tree resistance and beetle aggregation on bark beetle outbreaks. In this article we develop a new mathematical model for bark beetle outbreaks using an analogy with epidemiological models. Because the model operates on several distinct time scales, singular perturbation methods are used to simplify the model. The result is a dynamical system that tracks populations of uninfested and infested trees. A limiting case of the model is a discontinuous function of state variables, leading to solutions in the Filippov sense. The model assumes an extensive seed-bank so that tree recruitment is possible even if trees go extinct. Two scenarios are considered for immigration of new beetles. The first is a single tree stand with beetles immigrating from outside while the second considers two forest stands with beetle dispersal between them. For the seed-bank driven recruitment rate, when beetle immigration is low, the forest stand recovers to a beetle-free state. At high beetle immigration rates beetle populations approach an endemic equilibrium state. At intermediate immigration rates, the model predicts bistability as the forest can be in either of the two equilibrium states: a healthy forest, or a forest with an endemic beetle population. The model bistability leads to hysteresis. Interactions between two stands show how a less resistant stand of trees may provide an initial toe-hold for the invasion, which later leads to a regional beetle outbreak in the resistant stand.

Tree mortality from drought, insects, and their interactions in a changing climate.

Climate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects - bark beetles and defoliators - which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree-insect interactions will better inform projections of forest ecosystem responses to climate change.

Integrating models to investigate critical phenological overlaps in complex ecological interactions: the mountain pine beetle-fungus symbiosis.

The fates of individual species are often tied to synchronization of phenology, however, few methods have been developed for integrating phenological models involving linked species. In this paper, we focus on mountain pine beetle (MPB, Dendroctonus ponderosae) and its two obligate mutualistic fungi, Grosmannia clavigera and Ophiostoma montium. Growth rates of all three partners are driven by temperature, and their idiosyncratic responses affect interactions at important life stage junctures. One critical phase for MPB-fungus symbiosis occurs just before dispersal of teneral (new) adult beetles, when fungi are acquired and transported in specialized structures (mycangia). Before dispersal, fungi must capture sufficient spatial resources within the tree to ensure contact with teneral adults and get packed into mycangia. Mycangial packing occurs at an unknown time during teneral feeding. We adapt thermal models predicting fungal growth and beetle development to predict overlap between the competing fungi and MPB teneral adult feeding windows and emergence. We consider a spectrum of mycangial packing strategies and describe them in terms of explicit functions with unknown parameters. Rates of growth are fixed by laboratory data, the unknown parameters describing various packing strategies, as well as the degree to which mycangial growth is slowed in woody tissues as compared to agar, are determined by maximum likelihood and two years of field observations. At the field location used, the most likely fungus acquisition strategy for MPB was packing mycangia just prior to emergence. Estimated model parameters suggested large differences in the relative growth rates of the two fungi in trees at the study site, with the most likely model estimating that G. clavigera grew approximately twenty-five times faster than O. montium under the bark, which is completely unexpected in comparison with observed fungal growth on agar.

Mountain pine beetle seasonal timing and constraints to bivoltinism.

Mountain pine beetle tree colonization typically occurs in July and August, with completion of a generation one (univoltinism) or two (semivoltinism) years later. In a 2012 publication, Mitton and Ferrenberg suggested that climate change resulted in an unprecedented generation between June and September (a summer generation), with a concomitant shift to two generations in one year (bivoltinism). Although summer generations are not uncommon in this species, completion of a second generation across winter, between September and June, would be required for bivoltinism, a phenomenon not previously observed. Mitton and Ferrenberg showed that a summer generation can occur, but they failed to adequately track cohorts and provided no compelling evidence for bivoltinism. We demonstrate that a winter generation-and hence bivoltinism-would have been physiologically impossible at the high-elevation site used in Mitton and Ferrenberg due to lower thermal developmental thresholds. The mountain pine beetle is indeed being influenced by climate change. To address the challenges of future population outbreaks of this significant tree mortality agent, however, it is imperative to consider evolved, thermally dependent traits that serve to maintain seasonality.

Evidence for an adult summer diapause in mountain pine beetle (Coleoptera: Curculionidae) that varies geographically and among haplogroups.

Identifying dormancy traits is important for predicting insect population success, particularly in a changing climate that could disrupt evolved traits. The mountain pine beetle (Dendroctonus ponderosae Hopkins) is native to North America, is responsible for millions of acres of tree mortality, and is expanding northward in Canada. Research has identified thermal traits important to epidemic-phase ecology that vary among populations. Genomic research identified 3 mountain pine beetle haplogroups representing Pleistocene glacial refugia. Significant variation in generation timing aligning with the haplogroups has been observed. The adult stage was previously identified as the likely cause of differences among populations, although the mechanism(s) remain unclear. We tested for an adult summer diapause that varies among populations from 2 haplogroups, southern Colorado (CO) (central haplogroup) and southern Idaho (ID) (eastern haplogroup) using respirometry and reproduction experiments. Warm temperatures (25 °C) resulted in reduced respiration rates of central haplogroup mountain pine beetle compared to a cool temperature treatment (15 °C), whereas respiration of the eastern haplogroup did not differ between the treatments. Mated pairs of central haplogroup mountain pine beetle reared/held at 15 °C were more likely to be classified with a higher reproductive success rating compared to pairs reared/held at 25 °C. These results support a facultative summer adult diapause in southern CO central haplogroup mountain pine beetle. Manifestation of this diapause was low/absent among adults from the northerly ID location. This diapause likely serves to maintain univoltinism shown to be important for mountain pine beetle epidemic-phase ecology. The variation occurring among haplogroups highlights the long-term, evolved processes driving local adaptations in mountain pine beetle.

Seasonality of forest insects: why diapause matters.

Insects have major impacts on forest ecosystems, from herbivory and soil-nutrient cycling to killing trees at a large scale. Forest insects from temperate, tropical, and subtropical regions have evolved strategies to respond to seasonality; for example, by entering diapause, to mitigate adversity and to synchronize lifecycles with favorable periods. Here, we show that distinct functional groups of forest insects; that is, canopy dwellers, trunk-associated species, and soil/litter-inhabiting insects, express a variety of diapause strategies, but do not show systematic differences in diapause strategy depending on functional group. Due to the overall similarities in diapause strategies, we can better estimate the impacts of anthropogenic change on forest insect populations and, consequently, on key ecosystems.

Lignin concentrations in phloem and outer bark are not associated with resistance to mountain pine beetle among high elevation pines.

A key component in understanding plant-insect interactions is the nature of host defenses. Research on defense traits among Pinus species has focused on specialized metabolites and axial resin ducts, but the role of lignin in defense within diverse systems is unclear. We investigated lignin levels in the outer bark and phloem of P. longaeva, P. balfouriana, and P. flexilis; tree species growing at high elevations in the western United States known to differ in susceptibility to mountain pine beetle (Dendroctonus ponderosae; MPB). Pinus longaeva and P. balfouriana are attacked by MPB less frequently than P. flexilis, and MPB brood production in P. longaeva is limited. Because greater lignification of feeding tissues has been shown to provide defense against bark beetles in related genera, such as Picea, we hypothesized that P. longaeva and P. balfouriana would have greater lignin concentrations than P. flexilis. Contrary to expectations, we found that the more MPB-susceptible P. flexilis had greater phloem lignin levels than the less susceptible P. longaeva and P. balfouriana. No differences in outer bark lignin levels among the species were found. We conclude that lignification in Pinus phloem and outer bark is likely not adaptive as a physical defense against MPB.

Oral Anticoagulation Use in High-Risk Patients Is Improved by Elimination of False-Positive and Inactive Atrial Fibrillation Cases.

BACKGROUND: Multiple registries have reported that >40% of high-risk atrial fibrillation patients are not taking oral anticoagulants. The purpose of our study was to determine the presence or absence of active atrial fibrillation and CHA2DS2-VASc (Congestive heart failure, Hypertension, Age ≥75 y, Diabetes mellitus, prior Stroke [or transient ischemic attack or thromboembolism], Vascular disease, Age 65-74 y, Sex category) risk factors to accurately identify high-risk atrial fibrillation (CHA2DS2-VASc ≥2) patients requiring oral anticoagulants and the magnitude of the anticoagulant treatment gap. METHODS: We retrospectively adjudicated 6514 patients with atrial fibrillation documented by at least one of: billing diagnosis, electronic medical record encounter diagnosis, electronic medical record problem list, or electrocardiogram interpretation. RESULTS: After review, 4555/6514 (69.9%) had active atrial fibrillation, while 1201 had no documented history of atrial fibrillation and 758 had a history of atrial fibrillation that was no longer active. After removing the 1201 patients without a confirmed atrial fibrillation diagnosis, oral anticoagulant use in high-risk patients increased to 71.1% (P < .0001 compared with 62.9% at baseline). Oral anticoagulant use increased to 79.7% when the 758 inactive atrial fibrillation patients were also eliminated from the analysis (P < .0001 compared with baseline). In the active high-risk atrial fibrillation group, there was no significant difference in the use of oral anticoagulants between men (80.7%) and women (78.8%) with a CHA2DS2-VASc ≥2, or in women with a CHA2DS2-VASc ≥3 (79.9%). CONCLUSIONS: Current registries and health system health records with unadjudicated diagnoses over-report the number of high-risk atrial fibrillation patients not taking oral anticoagulants. Expert adjudication identifies a smaller treatment gap than previously described.

The Fire and Tree Mortality Database, for empirical modeling of individual tree mortality after fire.

Wildland fires have a multitude of ecological effects in forests, woodlands, and savannas across the globe. A major focus of past research has been on tree mortality from fire, as trees provide a vast range of biological services. We assembled a database of individual-tree records from prescribed fires and wildfires in the United States. The Fire and Tree Mortality (FTM) database includes records from 164,293 individual trees with records of fire injury (crown scorch, bole char, etc.), tree diameter, and either mortality or top-kill up to ten years post-fire. Data span 142 species and 62 genera, from 409 fires occurring from 1981-2016. Additional variables such as insect attack are included when available. The FTM database can be used to evaluate individual fire-caused mortality models for pre-fire planning and post-fire decision support, to develop improved models, and to explore general patterns of individual fire-induced tree death. The database can also be used to identify knowledge gaps that could be addressed in future research.

Evidence for a Prepupal Diapause in the Mountain Pine Beetle (Dendroctonus ponderosae).

Dormancy strategies, including diapause and quiescence, enable insects to evade adverse conditions and ensure seasonally appropriate life stages. A mechanistic understanding of a species' dormancy is necessary to predict population response in a changing climate. Climate change is influencing distribution patterns and population success of many species, including Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae: Scolytinae), the most important mortality agent of pines in western North America. Diapause is considered absent in D. ponderosae, and quiescence in the final larval stage prior to pupation (i.e., prepupal) is considered the main dormancy strategy. We evaluated if a facultative diapause in the prepupal stage, rather than a pupation threshold ~15°C (i.e., quiescence), could describe pupation patterns in two latitudinally separated D. ponderosae populations in the western United States. We hypothesized that if pupation occurs at lower temperatures than previously described, and if significant prepupal developmental delays occur, diapause is a likely physiological mechanism. Although there was considerable variation within and between populations, pupation occurred below the previously established threshold suggesting a prepupal facultative diapause that is induced when late instars experience cool temperatures. Individuals that pupated at temperatures below 15°C also had developmental delays, relative to development at warmer temperatures, consistent with diapause development. Pupation patterns differed between populations wherein diapause was induced at cooler temperatures and diapause development was shorter in southern compared with northern D. ponderosae. Recognition of a facultative diapause that varies among and between populations is critical for making predictions about future population response and range expansion in a changing climate.

Low offspring survival in mountain pine beetle infesting the resistant Great Basin bristlecone pine supports the preference-performance hypothesis.

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.

Modeling cold tolerance in the mountain pine beetle, Dendroctonus ponderosae.

Cold-induced mortality is a key factor driving mountain pine beetle, Dendroctonus ponderosae, population dynamics. In this species, the supercooling point (SCP) is representative of mortality induced by acute cold exposure. Mountain pine beetle SCP and associated cold-induced mortality fluctuate throughout a generation, with the highest SCPs prior to and following winter. Using observed SCPs of field-collected D. ponderosae larvae throughout the developmental season and associated phloem temperatures, we developed a mechanistic model that describes the SCP distribution of a population as a function of daily changes in the temperature-dependent processes leading to gain and loss of cold tolerance. It is based on the changing proportion of individuals in three states: (1) a non cold-hardened, feeding state, (2) an intermediate state in which insects have ceased feeding, voided their gut content and eliminated as many ice-nucleating agents as possible from the body, and (3) a fully cold-hardened state where insects have accumulated a maximum concentration of cryoprotectants (e.g. glycerol). Shifts in the proportion of individuals in each state occur in response to the driving variables influencing the opposite rates of gain and loss of cold hardening. The level of cold-induced mortality predicted by the model and its relation to extreme winter temperature is in good agreement with a range of field and laboratory observations. Our model predicts that cold tolerance of D. ponderosae varies within a season, among seasons, and among geographic locations depending on local climate. This variability is an emergent property of the model, and has important implications for understanding the insect's response to seasonal fluctuations in temperature, as well as population response to climate change. Because cold-induced mortality is but one of several major influences of climate on D. ponderosae population dynamics, we suggest that this model be integrated with others simulating the insect's biology.