Relative water content consistently predicts drought mortality risk in seedling populations with different morphology, physiology and times to death.

Predicted increases in forest drought mortality highlight the need for predictors of incipient drought-induced mortality (DIM) risk that enable proactive large-scale management. Such predictors should be consistent across plants with varying morphology and physiology. Because of their integrative nature, indicators of water status are promising candidates for real-time monitoring of DIM, particularly if they standardize morphological differences among plants. We assessed the extent to which differences in morphology and physiology between Pinus ponderosa populations influence time to mortality and the predictive power of key indicators of DIM risk. Time to incipient mortality differed between populations but occurred at the same relative water content (RWC) and water potential (WP). RWC and WP were accurate predictors of drought mortality risk. These results highlight that variables related to water status capture critical thresholds during DIM and the associated dehydration processes. Both WP and RWC are promising candidates for large-scale assessments of DIM risk. RWC is of special interest because it allows comparisons across different morphologies and can be remotely sensed. Our results offer promise for real-time landscape-level monitoring of DIM and its global impacts in the near term.

Bimodal cambial activity and false-ring formation in conifers under a monsoon climate.

Tracking wood formation in semiarid regions during the seasonal march of precipitation extremes has two important applications. It can provide (i) insight into the adaptive capacities of trees to drought and (ii) a basis for a richer interpretation of tree-ring data, assisting in a deeper understanding of past and current climate. In the southwestern USA, the anatomical signature of seasonally bimodal precipitation is the 'false ring'-a band of latewood-like cells in the earlywood. These occur when a particularly deep drought during the early growing season ends abruptly with timely, mid-growing season monsoonal rains. Such conditions presented in southern Arizona in 2014, enabling us to explore false-ring formation in ponderosa pine (Pinus ponderosa Lawson and C. Lawson) and Douglas-fir (Pseudotsuga menziesii Mirb. Franco) in mixed-conifer forest at 2573 m above sea level. We ask: what were the cell-by-cell timings and durations in the phases of wood cell development in 2014? How do these seasonal patterns relate to strongly fluctuating environmental conditions during the growing season? We took weekly microcores from March through November from six ponderosa pine and seven Douglas-fir trees at a well-instrumented flux tower site. Thin sections were prepared, and we counted cells in cambial, expansion, cell wall thickening and mature phases. For ponderosa pine trees forming a false ring, the first impact of intensifying seasonal drought was seen in the enlarging phase and then, almost a month later, in cambial activity. In this species, recovery from drought was associated with recovery first in cambial activity, followed by cell enlargement. This timing raised the possibility that cell division may be affected by atmospheric moisture increases before soil recharge. In both species, the last false-ring cells matured during the summer rainy season. Bimodal cambial activity coincident with moisture availability was observed in both species, whether or not they formed a false ring. This deeper knowledge of the precise timing of both developmental and environmental events should help define mechanistic connections among these factors in creating bimodal growth patterns.

Still standing: Recent patterns of post-fire conifer refugia in ponderosa pine-dominated forests of the Colorado Front Range.

Forested fire refugia (trees that survive fires) are important disturbance legacies that provide seed sources for post-fire regeneration. Conifer regeneration has been limited following some recent western fires, particularly in ponderosa pine (Pinus ponderosa) forests. However, the extent, characteristics, and predictability of ponderosa pine fire refugia are largely unknown. Within 23 fires in ponderosa pine-dominated forests of the Colorado Front Range (1996-2013), we evaluated the spatial characteristics and predictability of refugia: first using Monitoring Trends in Burn Severity (MTBS) burn severity metrics, then using landscape variables (topography, weather, anthropogenic factors, and pre-fire forest cover). Using 1-m resolution aerial imagery, we created a binary variable of post-fire conifer presence ('Conifer Refugia') and absence ('Conifer Absence') within 30-m grid cells. We found that maximum patch size of Conifer Absence was positively correlated with fire size, and 38% of the burned area was ≥ 50m from a conifer seed source, revealing a management challenge as fire sizes increase with warming further limiting conifer recovery. In predicting Conifer Refugia with two MTBS-produced databases, thematic burn severity classes (TBSC) and continuous Relative differenced Normalized Burn Ratio (RdNBR) values, Conifer Absence was high in previously forested areas of Low and Moderate burn severity classes in TBSC. RdNBR more accurately identified post-fire conifer survivorship. In predicting Conifer Refugia with landscape variables, Conifer Refugia were less likely during burn days with high maximum temperatures: while Conifer Refugia were more likely on moister soils and closer to higher order streams, homes, and roads; and on less rugged, valley topography. Importantly, pre-fire forest canopy cover was not strongly associated with Conifer Refugia. This study further informs forest management by mapping post-fire patches lacking conifer seed sources, validating the use of RdNBR for fire refugia, and detecting abiotic and topographic variables that may promote conifer refugia.

Conflicting functional effects of xylem pit structure relate to the growth-longevity trade-off in a conifer species.

Consistent with a ubiquitous life history trade-off, trees exhibit a negative relationship between growth and longevity both among and within species. However, the mechanistic basis of this life history trade-off is not well understood. In addition to resource allocation conflicts among multiple traits, functional conflicts arising from individual morphological traits may also contribute to life history trade-offs. We hypothesized that conflicting functional effects of xylem structural traits contribute to the growth-longevity trade-off in trees. We tested this hypothesis by examining the extent to which xylem morphological traits (i.e., wood density, tracheid diameters, and pit structure) relate to growth rates and longevity in two natural populations of the conifer species Pinus ponderosa Hydraulic constraints arise as trees grow larger and xylem anatomical traits adjust to compensate. We disentangled the effects of size through ontogeny in individual trees and growth rates among trees on xylem traits by sampling each tree at multiple trunk diameters. We found that the oldest trees had slower lifetime growth rates compared with younger trees in the studied populations, indicating a growth-longevity trade-off. We further provide evidence that a single xylem trait, pit structure, with conflicting effects on xylem function (hydraulic safety and efficiency) relates to the growth-longevity trade-off in a conifer species. This study highlights that, in addition to trade-offs among multiple traits, functional constraints based on individual morphological traits like that of pit structure provide mechanistic insight into how and when life history trade-offs arise.

Wildfires and climate change push low-elevation forests across a critical climate threshold for tree regeneration.

Climate change is increasing fire activity in the western United States, which has the potential to accelerate climate-induced shifts in vegetation communities. Wildfire can catalyze vegetation change by killing adult trees that could otherwise persist in climate conditions no longer suitable for seedling establishment and survival. Recently documented declines in postfire conifer recruitment in the western United States may be an example of this phenomenon. However, the role of annual climate variation and its interaction with long-term climate trends in driving these changes is poorly resolved. Here we examine the relationship between annual climate and postfire tree regeneration of two dominant, low-elevation conifers (ponderosa pine and Douglas-fir) using annually resolved establishment dates from 2,935 destructively sampled trees from 33 wildfires across four regions in the western United States. We show that regeneration had a nonlinear response to annual climate conditions, with distinct thresholds for recruitment based on vapor pressure deficit, soil moisture, and maximum surface temperature. At dry sites across our study region, seasonal to annual climate conditions over the past 20 years have crossed these thresholds, such that conditions have become increasingly unsuitable for regeneration. High fire severity and low seed availability further reduced the probability of postfire regeneration. Together, our results demonstrate that climate change combined with high severity fire is leading to increasingly fewer opportunities for seedlings to establish after wildfires and may lead to ecosystem transitions in low-elevation ponderosa pine and Douglas-fir forests across the western United States.

Carbon and oxygen isotope fractionations in tree rings reveal interactions between cambial phenology and seasonal climate.

We developed novel approaches for using the isotope composition of tree-ring subdivisions to study seasonal dynamics in tree-climate relations. Across a 30-year time series, the δ13 C and δ18 O values of the earlywood (EW) cellulose in the annual rings of Pinus ponderosa reflected relatively high intrinsic water-use efficiencies and high evaporative fractionation of 18 O/16 O, respectively, compared with the false latewood (FLW), summerwood (SW), and latewood (LW) subdivisions. This result is counterintuitive, given the spring origins of the EW source water and midsummer origins of the FLW, SW, and LW. With the use of the Craig-Gordon (CG), isotope-climate model revealed that the isotope ratios in all of the ring subdivision are explained by the existence of seasonal lags, lasting several weeks, between the initial formation of tracheids and the production of cellulosic secondary cell walls during maturation. In contrast to some past studies, modification of the CG model according to conventional methods to account for mixing of needle water between fractionated and nonfractionated sources did not improve the accuracy of predictions. Our results reveal new potential in the use of tree-ring isotopes to reconstruct past intra-annual tree-climate relations if lags in cambial phenology are reconciled with isotope ratio observations and included in theoretical treatments.

Moisture-driven xylogenesis in Pinus ponderosa from a Mojave Desert mountain reveals high phenological plasticity.

Future seasonal dynamics of wood formation in hyperarid environments are still unclear. Although temperature-driven extension of the growing season and increased forest productivity are expected for boreal and temperate biomes under global warming, a similar trend remains questionable in water-limited regions. We monitored cambial activity in a montane stand of ponderosa pine (Pinus ponderosa) from the Mojave Desert for 2 consecutive years (2015-2016) showing opposite-sign anomalies between warm- and cold-season precipitation. After the wet winter/spring of 2016, xylogenesis started 2 months earlier compared to 2015, characterized by abundant monsoonal (July-August) rainfall and hyperarid spring. Tree size did not influence the onset and ending of wood formation, highlighting a predominant climatic control over xylem phenological processes. Moisture conditions in the previous month, in particular soil water content and dew point, were the main drivers of cambial phenology. Latewood formation started roughly at the same time in both years; however, monsoonal precipitation triggered the formation of more false rings and density fluctuations in 2015. Because of uncertainties in future precipitation patterns simulated by global change models for the Southwestern United States, the dependency of P. ponderosa on seasonal moisture implies a greater conservation challenge than for species that respond mostly to temperature conditions.

Single-provenance mature conifers show higher non-structural carbohydrate storage and reduced growth in a drier location.

Since growth is more sensitive to drought than photosynthesis, trees inhabiting dry regions are expected to exhibit higher carbohydrate storage and less growth than their conspecifics from more humid regions. However, the same pattern can be the result of different genotypes inhabiting contrasting humidity conditions. To test if reduced growth and high carbohydrate storage are environmentally driven by drought, we examined the growth and non-structural carbohydrate (NSC) concentrations in single-provenance stands of mature trees of Pinus contorta Douglas and Pinus ponderosa Douglas ex C. Lawson planted at contrasting humidity conditions (900 versus 300 mm of annual precipitation) in Patagonia, Chile. Individual tree growth was measured for each species and at each location as mean basal area increment of the last 10 years (BAI10), annual shoot elongation for the period 2011-14, and needle length for 2013 and 2014 cohorts. Additionally, needle, branch, stem sapwood and roots were collected from each sampled tree to determine soluble sugars, starch and total NSC concentrations. The two species showed lower mean BAI10 and 2013 needle length in the dry site; P. ponderosa also had lower annual shoot extension for 2011 and 2014, and lower 2014 needle length, in the dry than in the mesic site. By contrast, NSC concentrations of all woody tissues for both species were either similar or higher in the dry site when compared with the mesic site. Patterns of starch and sugars were substantially different: starch concentrations were similar between sites except for roots of P. ponderosa, which were higher in the dry site, while sugar concentrations of all woody tissues in both species were higher in the dry site. Overall, our study provides evidence that reduced growth along with carbon (C) accumulation is an environmentally driven response to drought. Furthermore, the significant accumulation of low-molecular weight sugars in the dry site is compatible with a prioritized C allocation for osmoregulation. However, since this accumulation did not come at the expense of reduced starch, it is unlikely that growth was limited by C supply in the dry site.

Sapwood Stored Resources Decline in Whitebark and Lodgepole Pines Attacked by Mountain Pine Beetles (Coleoptera: Curculionidae).

Recent outbreaks of forest insects have been directly linked to climate change-induced warming and drought, but effects of tree stored resources on insects have received less attention. We asked whether tree stored resources changed following mountain pine beetle (Dendroctonus ponderosae Hopkins) attack and whether they affected beetle development. We compared initial concentrations of stored resources in the sapwood of whitebark pine (Pinus albicaulis Engelmann) and lodgepole pine (Pinus contorta Douglas ex. Louden) with resource concentrations one year later, in trees that were naturally attacked by beetles and trees that remained unattacked. Beetles did not select host trees based on sapwood resources-there were no consistent a priori differences between attacked versus unattacked trees-but concentrations of nonstructural carbohydrate (NSC), lipids, and phosphorus declined in attacked trees, relative to initial concentrations and unattacked trees. Whitebark pine experienced greater resource declines than lodgepole pine; however, sapwood resources were not correlated with beetle success in either species. Experimental manipulation confirmed that the negative effect of beetles on sapwood and phloem NSC was not due to girdling. Instead, changes in sapwood resources were related to the percentage of sapwood with fungal blue-stain. Overall, mountain pine beetle attack affected sapwood resources, but sapwood resources did not contribute directly to beetle success; instead, sapwood resources may support colonization by beetle-vectored fungi that potentially accelerate tree mortality. Closer attention to stored resource dynamics will improve our understanding of the interaction between mountain pine beetles, fungi, and host trees, an issue that is relevant to our understanding of insect range expansion under climate change.

Multiscale perspectives of fire, climate and humans in western North America and the Jemez Mountains, USA.

Interannual climate variations have been important drivers of wildfire occurrence in ponderosa pine forests across western North America for at least 400 years, but at finer scales of mountain ranges and landscapes human land uses sometimes over-rode climate influences. We reconstruct and analyse effects of high human population densities in forests of the Jemez Mountains, New Mexico from ca 1300 CE to Present. Prior to the 1680 Pueblo Revolt, human land uses reduced the occurrence of widespread fires while simultaneously adding more ignitions resulting in many small-extent fires. During the 18th and 19th centuries, wet/dry oscillations and their effects on fuels dynamics controlled widespread fire occurrence. In the late 19th century, intensive livestock grazing disrupted fuels continuity and fire spread and then active fire suppression maintained the absence of widespread surface fires during most of the 20th century. The abundance and continuity of fuels is the most important controlling variable in fire regimes of these semi-arid forests. Reduction of widespread fires owing to reduction of fuel continuity emerges as a hallmark of extensive human impacts on past forests and fire regimes.This article is part of the themed issue 'The interaction of fire and mankind'.

Drought stress limits the geographic ranges of two tree species via different physiological mechanisms.

Range shifts are among the most ubiquitous ecological responses to anthropogenic climate change and have large consequences for ecosystems. Unfortunately, the ecophysiological forces that constrain range boundaries are poorly understood, making it difficult to mechanistically project range shifts. To explore the physiological mechanisms by which drought stress controls dry range boundaries in trees, we quantified elevational variation in drought tolerance and in drought avoidance-related functional traits of a widespread gymnosperm (ponderosa pine - Pinus ponderosa) and angiosperm (trembling aspen - Populus tremuloides) tree species in the southwestern USA. Specifically, we quantified tree-to-tree variation in growth, water stress (predawn and midday xylem tension), drought avoidance traits (branch conductivity, leaf/needle size, tree height, leaf area-to-sapwood area ratio), and drought tolerance traits (xylem resistance to embolism, hydraulic safety margin, wood density) at the range margins and range center of each species. Although water stress increased and growth declined strongly at lower range margins of both species, ponderosa pine and aspen showed contrasting patterns of clinal trait variation. Trembling aspen increased its drought tolerance at its dry range edge by growing stronger but more carbon dense branch and leaf tissues, implying an increased cost of growth at its range boundary. By contrast, ponderosa pine showed little elevational variation in drought-related traits but avoided drought stress at low elevations by limiting transpiration through stomatal closure, such that its dry range boundary is associated with limited carbon assimilation even in average climatic conditions. Thus, the same climatic factor (drought) may drive range boundaries through different physiological mechanisms - a result that has important implications for process-based modeling approaches to tree biogeography. Further, we show that comparing intraspecific patterns of trait variation across ranges, something rarely done in a range-limit context, helps elucidate a mechanistic understanding of range constraints.

Ponderosa pine resin defenses and growth: metrics matter.

Bark beetles (Coleoptera: Curculionidae, Scolytinae) cause widespread tree mortality in coniferous forests worldwide. Constitutive and induced host defenses are important factors in an individual tree's ability to survive an attack and in bottom-up regulation of bark beetle population dynamics, yet quantifying defense levels is often difficult. For example, in Pinus spp., resin flow is important for resistance to bark beetles but is extremely variable among individuals and within a season. While resin is produced and stored in resin ducts, the specific resin duct metrics that best correlate with resin flow remain unclear. The ability and timing of some pine species to produce induced resin is also not well understood. We investigated (i) the relationships between ponderosa pine (Pinus ponderosa Lawson & C. Lawson) resin flow and axial resin duct characteristics, tree growth and physiological variables, and (ii) if mechanical wounding induces ponderosa pine resin flow and resin ducts in the absence of bark beetles. Resin flow increased later in the growing season under moderate water stress and was highest in faster growing trees. The best predictors of resin flow were nonstandardized measures of resin ducts, resin duct size and total resin duct area, both of which increased with tree growth. However, while faster growing trees tended to produce more resin, models of resin flow using only tree growth were not statistically significant. Further, the standardized measures of resin ducts, density and duct area relative to xylem area, decreased with tree growth rate, indicating that slower growing trees invested more in resin duct defenses per unit area of radial growth, despite a tendency to produce less resin overall. We also found that mechanical wounding induced ponderosa pine defenses, but this response was slow. Resin flow increased after 28 days, and resin duct production did not increase until the following year. These slow induced responses may allow unsuccessfully attacked or wounded trees to resist future bark beetle attacks. Forest management that encourages healthy, vigorously growing trees will also favor larger resin ducts, thereby conferring increased constitutive resistance to bark beetle attacks.

An integrated model of environmental effects on growth, carbohydrate balance, and mortality of Pinus ponderosa forests in the southern Rocky Mountains.

Climate-induced tree mortality is an increasing concern for forest managers around the world. We used a coupled hydrologic and ecosystem carbon cycling model to assess temperature and precipitation impacts on productivity and survival of ponderosa pine (Pinus ponderosa). Model predictions were evaluated using observations of productivity and survival for three ponderosa pine stands located across an 800 m elevation gradient in the southern Rocky Mountains, USA, during a 10-year period that ended in a severe drought and extensive tree mortality at the lowest elevation site. We demonstrate the utility of a relatively simple representation of declines in non-structural carbohydrate (NSC) as an approach for estimating patterns of ponderosa pine vulnerability to drought and the likelihood of survival along an elevation gradient. We assess the sensitivity of simulated net primary production, NSC storage dynamics, and mortality to site climate and soil characteristics as well as uncertainty in the allocation of carbon to the NSC pool. For a fairly wide set of assumptions, the model estimates captured elevational gradients and temporal patterns in growth and biomass. Model results that best predict mortality risk also yield productivity, leaf area, and biomass estimates that are qualitatively consistent with observations across the sites. Using this constrained set of parameters, we found that productivity and likelihood of survival were equally dependent on elevation-driven variation in temperature and precipitation. Our results demonstrate the potential for a coupled hydrology-ecosystem carbon cycling model that includes a simple model of NSC dynamics to predict drought-related mortality. Given that increases in temperature and in the frequency and severity of drought are predicted for a broad range of ponderosa pine and other western North America conifer forest habitats, the model potentially has broad utility for assessing ecosystem vulnerabilities.

Sensitivity of ring growth and carbon allocation to climatic variation vary within ponderosa pine trees.

Most dendrochronological studies focus on cores sampled from standard positions (main stem, breast height), yet vertical gradients in hydraulic constraints and priorities for carbon allocation may contribute to different growth sensitivities with position. Using cores taken from five positions (coarse roots, breast height, base of live crown, mid-crown branch and treetop), we investigated how radial growth sensitivity to climate over the period of 1895-2008 varies by position within 36 large ponderosa pines (Pinus ponderosa Dougl.) in northern Arizona. The climate parameters investigated were Palmer Drought Severity Index, water year and monsoon precipitation, maximum annual temperature, minimum annual temperature and average annual temperature. For each study tree, we generated Pearson correlation coefficients between ring width indices from each position and six climate parameters. We also investigated whether the number of missing rings differed among positions and bole heights. We found that tree density did not significantly influence climatic sensitivity to any of the climate parameters investigated at any of the sample positions. Results from three types of analyses suggest that climatic sensitivity of tree growth varied with position height: (i) correlations of radial growth and climate variables consistently increased with height; (ii) model strength based on Akaike's information criterion increased with height, where treetop growth consistently had the highest sensitivity and coarse roots the lowest sensitivity to each climatic parameter; and (iii) the correlation between bole ring width indices decreased with distance between positions. We speculate that increased sensitivity to climate at higher positions is related to hydraulic limitation because higher positions experience greater xylem tensions due to gravitational effects that render these positions more sensitive to climatic stresses. The low sensitivity of root growth to all climatic variables measured suggests that tree carbon allocation to coarse roots is independent of annual climate variability. The greater number of missing rings in branches highlights the fact that canopy development is a low priority for carbon allocation during poor growing conditions.

Paired comparison of water, energy and carbon exchanges over two young maritime pine stands (Pinus pinaster Ait.): effects of thinning and weeding in the early stage of tree growth.

The effects of management practices on energy, water and carbon exchanges were investigated in a young pine plantation in south-west France. In 2009-10, carbon dioxide (CO(2)), H(2)O and heat fluxes were monitored using the eddy covariance and sap flow techniques in a control plot (C) with a developed gorse layer, and an adjacent plot that was mechanically weeded and thinned (W). Despite large differences in the total leaf area index and canopy structure, the annual net radiation absorbed was only 4% lower in plot W. We showed that higher albedo in this plot was offset by lower emitted long-wave radiation. Annual evapotranspiration (ET) from plot W was 15% lower, due to lower rainfall interception and transpiration by the tree canopy, partly counterbalanced by the larger evaporation from both soil and regrowing weedy vegetation. The drainage belowground from plot W was larger by 113 mm annually. The seasonal variability of ET was driven by the dynamics of the soil and weed layers, which was more severely affected by drought in plot C. Conversely, the temporal changes in pine transpiration and stem diameter growth were synchronous between sites despite higher soil water content in the weeded plot. At the annual scale, both plots were carbon sinks, but thinning and weeding reduced the carbon uptake by 73%: annual carbon uptake was 243 and 65 g C m(-2) on plots C and W, respectively. Summer drought dramatically impacted the net ecosystem exchange: plot C became a carbon source as the gross primary production (GPP) severely decreased. However, plot W remained a carbon sink during drought, as a result of decreases in both GPP and ecosystem respiration (R(E)). In winter, both plots were carbon sources, plots C and W emitting 67.5 and 32.4 g C m(-2), respectively. Overall, this study highlighted the significant contribution of the gorse layer to mass and energy exchange in young pine plantations.

Climate-related trends in sapwood biophysical properties in two conifers: avoidance of hydraulic dysfunction through coordinated adjustments in xylem efficiency, safety and capacitance.

In the Pacific north-west, the Cascade Mountain Range blocks much of the precipitation and maritime influence of the Pacific Ocean, resulting in distinct climates east and west of the mountains. The current study aimed to investigate relationships between water storage and transport properties in populations of Douglas-fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) adapted to both climates. Sapwood thickness, capacitance, vulnerability to embolism, and axial and radial conductivity were measured on samples collected from trunks of mature trees. The sapwood of ponderosa pine was three to four times thicker than Douglas-fir. Radial conductivity was higher in west-side populations of both species, but axial conductivity was higher in the east-side populations and in Douglas-fir. Eastern populations of both species had sapwood that was more vulnerable to embolism than west-side populations. Sapwood capacitance was similar between species, but was about twice as great in east-side populations (580 kg m⁻³ MPa⁻¹) as in west-side populations (274 kg m⁻³ MPa⁻¹). Capacitance was positively correlated with both mean embolism pressure and axial conductivity across species and populations, suggesting that coordinated adjustments in xylem efficiency, safety and water storage capacity may serve to avoid embolism along a gradient of increasing aridity.

Masting in ponderosa pine: comparisons of pollen and seed over space and time.

Many plant species exhibit variable and synchronized reproduction, or masting, but less is known of the spatial scale of synchrony, effects of climate, or differences between patterns of pollen and seed production. We monitored pollen and seed cone production for seven Pinus ponderosa populations (607 trees) separated by up to 28 km and 1,350 m in elevation in Boulder County, Colorado, USA for periods of 4-31 years for a mean per site of 8.7 years for pollen and 12.1 for seed cone production. We also analyzed climate data and a published dataset on 21 years of seed production for an eighth population (Manitou) 100 km away. Individual trees showed high inter-annual variation in reproduction. Synchrony was high within populations, but quickly became asynchronous among populations with a combination of increasing distance and elevational difference. Inter-annual variation in temperature and precipitation had differing influences on seed production for Boulder County and Manitou. We speculate that geographically variable effects of climate on reproduction arise from environmental heterogeneity and population genetic differentiation, which in turn result in localized synchrony. Although individual pines produce pollen and seed, only one-third of the covariation within trees was shared. As compared to seed cones, pollen had lower inter-annual variation at the level of the individual tree and was more synchronous. However, pollen and seed production were similar with respect to inter-annual variation at the population level, spatial scales of synchrony and associations with climate. Our results show that strong masting can occur at a localized scale, and that reproductive patterns can differ between pollen and seed cone production in a hermaphroditic plant.

Carbon and water fluxes from ponderosa pine forests disturbed by wildfire and thinning.

Disturbances alter ecosystem carbon dynamics, often by reducing carbon uptake and stocks. We compared the impact of two types of disturbances that represent the most likely future conditions of currently dense ponderosa pine forests of the southwestern United States: (1) high-intensity fire and (2) thinning, designed to reduce fire intensity. High-severity fire had a larger impact on ecosystem carbon uptake and storage than thinning. Total ecosystem carbon was 42% lower at the intensely burned site, 10 years after burning, than at the undisturbed site. Eddy covariance measurements over two years showed that the burned site was a net annual source of carbon to the atmosphere whereas the undisturbed site was a sink. Net primary production (NPP), evapotranspiration (ET), and water use efficiency were lower at the burned site than at the undisturbed site. In contrast, thinning decreased total ecosystem carbon by 18%, and changed the site from a carbon sink to a source in the first posttreatment year. Thinning also decreased ET, reduced the limitation of drought on carbon uptake during summer, and did not change water use efficiency. Both disturbances reduced ecosystem carbon uptake by decreasing gross primary production (55% by burning, 30% by thinning) more than total ecosystem respiration (TER; 33-47% by burning, 18% by thinning), and increased the contribution of soil carbon dioxide efflux to TER. The relationship between TER and temperature was not affected by either disturbance. Efforts to accurately estimate regional carbon budgets should consider impacts on carbon dynamics of both large disturbances, such as high-intensity fire, and the partial disturbance of thinning that is often used to prevent intense burning. Our results show that thinned forests of ponderosa pine in the southwestern United States are a desirable alternative to intensively burned forests to maintain carbon stocks and primary production.

Elevated CO2 and O3 effects on fine-root survivorship in ponderosa pine mesocosms.

Atmospheric carbon dioxide (CO(2)) and ozone (O(3)) concentrations are rising, which may have opposing effects on tree C balance and allocation to fine roots. More information is needed on interactive CO(2) and O(3) effects on roots, particularly fine-root life span, a critical demographic parameter and determinant of soil C and N pools and cycling rates. We conducted a study in which ponderosa pine (Pinus ponderosa) seedlings were exposed to two levels of CO(2) and O(3) in sun-lit controlled-environment mesocosms for 3 years. Minirhizotrons were used to monitor individual fine roots in three soil horizons every 28 days. Proportional hazards regression was used to analyze effects of CO(2), O(3), diameter, depth, and season of root initiation on fine-root survivorship. More fine roots were produced in the elevated CO(2) treatment than in ambient CO(2). Elevated CO(2), increasing root diameter, and increasing root depth all significantly increased fine-root survivorship and median life span. Life span was slightly, but not significantly, lower in elevated O(3), and increased O(3) did not reduce the effect of elevated CO(2). Median life spans varied from 140 to 448 days depending on the season of root initiation. These results indicate the potential for elevated CO(2) to increase the number of fine roots and their residence time in the soil, which is also affected by root diameter, root depth, and phenology.