ABSTRACT
Serotiny is an adaptive trait that allows certain woody plants to persist in stand-replacing fire regimes. However, the mechanisms by which serotinous cones avoid seed necrosis and nonserotinous species persist in landscapes with short fire cycles and serotinous competitors remain poorly understood. To investigate whether ovulate cone traits that enhance seed survival differ between serotinous and nonserotinous species, we examined cone traits in 24 species within Pinaceae and Cupressaceae based on physical measurements and cone heating simulations using a computational fluid dynamics model. Fire-relevant cone traits were largely similar between cone types; those that differed (e.g. density and moisture) conferred little seed survival advantage under simulated fire. The most important traits influencing seed survival were cone size and seed depth within the cone, which was found to be an allometric function of cone mass for both cone types. Thus, nonserotinous cones should not suffer significantly greater seed necrosis than serotinous cones of equal size. Closed nonserotinous cones containing mature seeds may achieve substantial regeneration after fire if they are sufficiently large relative to fire duration and temperature. To our knowledge, this is the most comprehensive study of the effects of fire-relevant cone traits on conifer regeneration supported by physics-based fire simulation.
Subject(s)
Fires , Tracheophyta , Seeds , Phenotype , NecrosisABSTRACT
Pinus species dominate fire-prone ecosystems throughout the northern hemisphere. Their litter drive fires that control plant community flammability and multiple ecological processes. To better understand the patterns and mechanisms of pine flammability, we measured leaf characteristics (needle length and thickness) and conducted combustion experiments on litter from 31 species. We paired flammability results with bark accumulation data and used phylogenetic generalized least squares regression to examine relationships between physical traits and flammability. Pine flammability varied widely among pines: flame heights and fuel consumption varied three-fold, and flaming and smoldering durations varied three- to six-fold. Subgenus Pinus species were the most flammable and subgenus Strobus species had the lowest flammability. Needle length was the best predictor of flammability with a significant interaction with subgenus, suggesting that flammability of pines in subgenus Strobus was more affected by physical traits than pines in subgenus Pinus. Species in the subgenus Pinus that accumulated outer bark rapidly also had high flammability, while the relationship was not significant in subgenus Strobus. These results highlight the diverse patterns of flammability in North American pines and the complexity in the mechanisms causing differential flammability.
Subject(s)
Fires , Pinus , Ecosystem , Phylogeny , Plant BarkABSTRACT
Resin ducts in the secondary xylem of tree rings are a measure of a tree's defense capacity from insects and pathogens. Because resin ducts are permanently embedded within the xylem, retrospective analysis can be performed to quantify changes in defense over time and determine factors that contribute to this change, such as climate and disturbance. Here, we provide methods on how to measure axial resin ducts in secondary xylem. These methods provide the necessary protocols for consistent quantification of xylem resin ducts and terminology, which will also allow easier cross-comparison among studies in the future. We describe:â¢Steps to prepare tree cores for resin duct measurements.â¢Procedure to obtain image and measure individual resin ducts.â¢Software code to compile duct measurements into a complete chronology with both standardized and unstandardized resin duct metrics for further analyses.
ABSTRACT
Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.
Subject(s)
Droughts , Trees/growth & development , Adaptation, Physiological , Climate Change , Cycadopsida/growth & development , Ecology , Forests , Magnoliopsida/growth & development , Mortality , Soil/chemistry , Species Specificity , Stress, Physiological , Survival Analysis , Trees/classification , WaterABSTRACT
PREMISE: Dispersal capacity primarily determines the spatial establishment patterns that drive range expansions and contractions in tree species. Seedling establishment in Baker cypress (Hesperocyparis bakeri [(Jeps.) Bartel]) relies predominantly on fire events, due to its cone serotiny, shade intolerance, and small seeds that require the optimal conditions of fire-exposed, mineral soil seedbeds. METHODS: We quantified the density and spatial distribution of post-disturbance seedlings following the 2014 Eiler Fire in northern California and compared the observed recruitment to predictions from a mechanistic seed dispersal model. RESULTS: Postfire recruitment was dense, averaging 11 seedlings/m2 , and occurred primarily in the first year after fire. We estimated the mean descent velocity of the wingless seeds as ~4 m/s, the highest value reported for any putatively wind-dispersed tree species. The rapid seed descent contributed to markedly spatially constrained recruitment. Most seedlings (~81%) established within 5 m of the parent tree, and 94% established within 10 m. The maximum observed dispersal distance was 48.5 m; dispersal distance scaled linearly with canopy height. Distributions of modeled seed dispersal distance and observed seedling establishment in Baker cypress did not differ, demonstrating that wind disperses seeds a short distance because of the lack of a wing, and secondary dispersal appeared to be minimal at this recently burned site. CONCLUSIONS: If seed dispersal is solely reliant on wind, migration in response to rapid climate change will be impeded and potentially present difficulties in sustaining populations of this and other obligate seeder species with equally constrained dispersal.
Subject(s)
Cupressus , Seed Dispersal , Tracheophyta , Animals , California , Seedlings , Seeds , TreesABSTRACT
Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter-annual growth variability and a decrease in growth synchrony in the last â¼20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.
ABSTRACT
Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.
Subject(s)
Carbon/deficiency , Droughts , Plant Transpiration/physiology , Trees/physiology , Xylem/physiology , Climate Change , Cycadopsida/physiology , Magnoliopsida/physiology , Population Dynamics , Stress, PhysiologicalABSTRACT
Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.
Subject(s)
Coleoptera , Droughts , Trees/growth & development , Animals , Carbon , Stress, PhysiologicalABSTRACT
Bark beetles have recently killed billions of trees, yet conifer defenses are formidable and some trees resist attack. A primary anti-insect defense of pines is oleoresin from a system of resin ducts throughout the tree. Resin defense traits are heritable, and evidence suggests that resin duct characteristics are associated with resistance to insects. However, comparisons of resin ducts in trees killed by bark beetles to trees that resisted attack are unavailable. We compared vertical resin duct characteristics (number, density, and size) and growth rates from trees that were "resistant" (survived mass attack) versus "susceptible" (killed by attack) to bark beetles in lodgepole (Pinus contorta) and limber (Pinus flexilis) pines. Resistant trees of both species had significantly more resin ducts in recent growth than susceptible trees. Discriminant analysis (DA) correctly categorized 84% of lodgepole and 92% of limber pines as susceptible/resistant based on combinations of resin duct and growth characteristics from recent 5- through 20-year growth intervals. DA models using measures from only the most recent 5 years of growth correctly categorized 72 and 81% of lodgepole and limber pines, respectively. Comparing resistant to susceptible trees independent of species identity led to the correct categorization of 82% of trees based on factors from 5- to 20-year intervals, and 73% of trees using only resin duct counts from the most recent 5 years. We conclude that resin duct characteristics can be used to assess tree resistance to bark beetles across pine species, and offer a metric for management to enhance pest resistance.
Subject(s)
Coleoptera , Pinus/anatomy & histology , Resins, Plant , Animals , Colorado , Discriminant Analysis , Pinus/growth & development , Trees/anatomy & histology , Trees/growth & developmentABSTRACT
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.
Subject(s)
Carbon/metabolism , Climate Change , Pinus ponderosa/growth & development , Pinus ponderosa/metabolism , Trees/growth & development , Trees/metabolism , Analysis of Variance , Arizona , Linear ModelsABSTRACT
The relative importance of growth and defense to tree mortality during drought and bark beetle attacks is poorly understood. We addressed this issue by comparing growth and defense characteristics between 25 pairs of ponderosa pine (Pinus ponderosa) trees that survived and trees that died from drought-associated bark beetle attacks in forests of northern Arizona, USA. The three major findings of our research were: (1) xylem resin ducts in live trees were >10% larger (diameter), >25% denser (no. of resin ducts mm(-2)), and composed >50% more area per unit ring growth than dead trees; (2) measures of defense, such as resin duct production (no. of resin ducts year(-1)) and the proportion of xylem ring area to resin ducts, not growth, were the best model parameters of ponderosa pine mortality; and (3) most correlations between annual variation in growth and resin duct characteristics were positive suggesting that conditions conducive to growth also increase resin duct production. Our results suggest that trees that survive drought and subsequent bark beetle attacks invest more carbon in resin defense than trees that die, and that carbon allocation to resin ducts is a more important determinant of tree mortality than allocation to radial growth.