Long‐term increases in intrinsic water‐use efficiency do not lead to increased stem growth in a tropical monsoon forest in western Thailand

Rising atmospheric carbon dioxide [CO₂] can accelerate tree growth by stimulating photosynthesis and increasing intrinsic water‐use efficiency (iWUE). Little evidence exists, however, for the long‐term growth and gas‐exchange responses of mature trees in tropical forests to the combined effects of rising [CO₂] and other global changes such as warming. Using tree rings and stable isotopes of carbon and oxygen, we investigated long‐term trends in the iWUE and stem growth (basal area increment, BAI) of three canopy tree species in a tropical monsoon forest in western Thailand (Chukrasia tabularis, Melia azedarach, and Toona ciliata). To do this, we modelled the contribution of ontogenetic effects (tree diameter or age) and calendar year to variation in iWUE, oxygen isotopes, and BAI using mixed‐effects models. Although iWUE increased significantly with both tree diameter and calendar year in all species, BAI at a given tree diameter was lower in more recent years. For one species, C. tabularis, differences in crown dominance significantly influence stable isotopes and growth. Tree ring Δ¹⁸O increased with calendar year in all species, suggesting that increasing iWUE may have been driven by relatively greater reductions in stomatal conductance – leading to enrichment in Δ¹⁸O – than increases in photosynthetic capacity. Plausible explanations for the observed declines in growth include water stress resulting from rising temperatures and El Niño events, increased respiration, changes in allocation, or more likely, a combination of these factors.     

No evidence for consistent long‐term growth stimulation of 13 tropical tree species: results from tree‐ring analysis

The important role of tropical forests in the global carbon cycle makes it imperative to assess changes in their carbon dynamics for accurate projections of future climate–vegetation feedbacks. Forest monitoring studies conducted over the past decades have found evidence for both increasing and decreasing growth rates of tropical forest trees. The limited duration of these studies restrained analyses to decadal scales, and it is still unclear whether growth changes occurred over longer time scales, as would be expected if CO₂‐fertilization stimulated tree growth. Furthermore, studies have so far dealt with changes in biomass gain at forest‐stand level, but insights into species‐specific growth changes – that ultimately determine community‐level responses – are lacking. Here, we analyse species‐specific growth changes on a centennial scale, using growth data from tree‐ring analysis for 13 tree species (~1300 trees), from three sites distributed across the tropics. We used an established (regional curve standardization) and a new (size‐class isolation) growth‐trend detection method and explicitly assessed the influence of biases on the trend detection. In addition, we assessed whether aggregated trends were present within and across study sites. We found evidence for decreasing growth rates over time for 8–10 species, whereas increases were noted for two species and one showed no trend. Additionally, we found evidence for weak aggregated growth decreases at the site in Thailand and when analysing all sites simultaneously. The observed growth reductions suggest deteriorating growth conditions, perhaps due to warming. However, other causes cannot be excluded, such as recovery from large‐scale disturbances or changing forest dynamics. Our findings contrast growth patterns that would be expected if elevated CO₂ would stimulate tree growth. These results suggest that commonly assumed growth increases of tropical forests may not occur, which could lead to erroneous predictions of carbon dynamics of tropical forest under climate change.     

Tree rings provide no evidence of a CO2 fertilization effect in old‐growth subalpine forests of western Canada

Atmospheric CO₂ concentrations are now 1.7 times higher than the preindustrial values. Although photosynthetic rates are hypothesized to increase in response to rising atmospheric CO₂ concentrations, results from in situ experiments are inconsistent in supporting a CO₂ fertilization effect of tree growth. Tree‐ring data provide a historical record of tree‐level productivity that can be used to evaluate long‐term responses of tree growth. We use tree‐ring data from old‐growth, subalpine forests of western Canada that have not had a stand‐replacing disturbance for hundreds of years to determine if growth has increased over 19th and 20th centuries. Our sample consisted of 5,858 trees belonging to five species distributed over two sites in the coastal zone and two in the continental climate of the interior. We calculated annual increments in tree basal area, adjusted these increments for tree size and age, and tested whether there was a detectable temporal trend in tree growth over the 19th and 20th centuries. We found a similar pattern in 20th century growth trends among all species at all sites. Growth during the 19th century was mostly stable or increasing, with the exception of one of the coastal sites, where tree growth was slightly decreasing; whereas growth during the 20th century consistently decreased. The unexpected decrease in growth during the 20th century indicates that there was no CO₂ fertilization effect on photosynthesis. We compared the growth trends from our four sites to the trends simulated by seven Earth System Models, and saw that most of the models did not predict these growth declines. Overall, our results indicate that these old‐growth forests are unlikely to increase their carbon storage capacity in response to rising atmospheric CO₂, and thus are unlikely to contribute substantially to offsetting future carbon emissions.     

Differential anatomical responses to elevated CO2 in saplings of four hardwood species

To determine whether an elevated carbon dioxide concentration ([CO₂]) can induce changes in the wood structure and stem radial growth in forest trees, we investigated the anatomical features of conduit cells and cambial activity in 4‐year‐old saplings of four deciduous broadleaved tree species – two ring‐porous (Quercus mongolica and Kalopanax septemlobus) and two diffuse‐porous species (Betula maximowicziana and Acer mono) – grown for three growing seasons in a free‐air CO₂ enrichment system. Elevated [CO₂] had no effects on vessels, growth and physiological traits of Q. mongolica, whereas tree height, photosynthesis and vessel area tended to increase in K. septemlobus. No effects of [CO₂] on growth, physiological traits and vessels were seen in the two diffuse‐porous woods. Elevated [CO₂] increased larger vessels in all species, except B. maximowicziana and number of cambial cells in two ring‐porous species. Our results showed that the vessel anatomy and radial stem growth of Q. mongolica, B. maximowicziana and A. mono were not affected by elevated [CO₂], although vessel size frequency and cambial activity in Q. mongolica were altered. In contrast, changes in vessel anatomy and cambial activity were induced by elevated [CO₂] in K. septemlobus. The different responses to elevated [CO₂] suggest that the sensitivity of forest trees to CO₂ is species dependent.     

Increase in CO2 concentration could alter the response of Hedera helix to climate change

Increasing CO₂ concentration ([CO₂]) is likely to affect future species distributions, in interaction with other climate change drivers. However, current modeling approaches still seldom consider interactions between climatic factors and the importance of these interactions therefore remains mostly unexplored. Here, we combined dendrochronological and modeling approaches to study the interactive effects of increasing [CO₂] and temperature on the distribution of one of the main European liana species, Hedera helix. We combined a classical continent‐wide species distribution modeling approach with a case study using H. helix and Quercus cerris tree rings, where we explored the long‐term influence of a variety of climate drivers, including increasing [CO₂], and their interactions, on secondary growth. Finally, we explored how our findings could influence the model predictions. Climate‐only model predictions showed a small decrease in habitat suitability for H. helix in Europe; however, this was accompanied by a strong shift in the distribution toward the north and east. Our growth ring data suggested that H. helix can benefit from high [CO₂] under warm conditions, more than its tree hosts, which showed a weaker response to [CO₂] coupled with higher cavitation risk under high temperature. Increasing [CO₂] might therefore offset the negative effects of high temperatures on H. helix, and we illustrate how this might translate into maintenance of H. helix in warmer areas. Our results highlight the need to consider carbon fertilization and interactions between climate variables in ecological modeling. Combining dendrochronological analyses with spatial distribution modeling may provide opportunities to refine predictions of how climate change will affect species distributions.     

Long‐term tree fern dynamics linked to disturbance and shade tolerance

Question: Do New Zealand tree ferns have recognizable shade tolerance niches? Location: Lowland temperate rain forest of New Zealand (41°20′S, 174°58′E). Methods: Growth, death and recruitment of five tree fern species were estimated from a 38‐year record of stem heights, collected within a 2.25‐ha block of forest, and electron transport rates (ETR) of photosystem II of fronds were measured. Results: Two species of Cyathea were comparatively common (603 and 351 stems in total) and two were comparatively rare (155 and 17 stems in total) on the site. The common species had lower rates of growth, recruitment and mortality than the rare species, had skewed age distributions typical of shade‐tolerant species and were probably recruited soon after a catastrophic earthquake in 1855. The two rare species were failing to recruit under closed forests; their age distributions indicated that all had regenerated long after the earthquake. ETR were higher for faster‐growing than for the shade‐tolerant species. A tree fern that regenerates vegetatively from aerial buds, Dicksonia squarrosa, was common on the site (361 stems in total). Its age distribution suggested it was relatively shade tolerant, but its mortality and recruitment rates were much higher than those of the two shade‐tolerating Cyathea species, suggesting that this multi‐stemmed species functions differently from the monopodial Cyathea species. Conclusions: New Zealand Cyathea tree ferns occupy distinct niches along a shade tolerance spectrum and their relative abundances are strongly influenced by disturbance history. The study provides evidence that tree fern species differ strongly in their responses to canopy disturbance and are not ecologically equivalent.     

Tree growth variation in the tropical forest: understanding effects of temperature,rainfall and CO2

Tropical forest responses to climatic variability have important consequences for global carbon cycling, but are poorly understood. As empirical, correlative studies cannot disentangle the interactive effects of climatic variables on tree growth, we used a tree growth model (IBTREE) to unravel the climate effects on different physiological pathways and in turn on stem growth variation. We parameterized the model for canopy trees of Toona ciliata (Meliaceae) from a Thai monsoon forest and compared predicted and measured variation from a tree‐ring study over a 30‐year period. We used historical climatic variation of minimum and maximum day temperature, precipitation and carbon dioxide (CO₂) in different combinations to estimate the contribution of each climate factor in explaining the inter‐annual variation in stem growth. Running the model with only variation in maximum temperature and rainfall yielded stem growth patterns that explained almost 70% of the observed inter‐annual variation in stem growth. Our results show that maximum temperature had a strong negative effect on the stem growth by increasing respiration, reducing stomatal conductance and thus mitigating a higher transpiration demand, and – to a lesser extent – by directly reducing photosynthesis. Although stem growth was rather weakly sensitive to rain, stem growth variation responded strongly and positively to rainfall variation owing to the strong inter‐annual fluctuations in rainfall. Minimum temperature and atmospheric CO₂ concentration did not significantly contribute to explaining the inter‐annual variation in stem growth. Our innovative approach – combining a simulation model with historical data on tree‐ring growth and climate – allowed disentangling the effects of strongly correlated climate variables on growth through different physiological pathways. Similar studies on different species and in different forest types are needed to further improve our understanding of the sensitivity of tropical tree growth to climatic variability and change.     

Predicting tree biomass growth in the temperate–boreal ecotone: Is tree size,age, competition,or climate response most important?

As global temperatures rise, variation in annual climate is also changing, with unknown consequences for forest biomes. Growing forests have the ability to capture atmospheric CO₂ and thereby slow rising CO₂ concentrations. Forests’ ongoing ability to sequester C depends on how tree communities respond to changes in climate variation. Much of what we know about tree and forest response to climate variation comes from tree‐ring records. Yet typical tree‐ring datasets and models do not capture the diversity of climate responses that exist within and among trees and species. We address this issue using a model that estimates individual tree response to climate variables while accounting for variation in individuals’ size, age, competitive status, and spatially structured latent covariates. Our model allows for inference about variance within and among species. We quantify how variables influence aboveground biomass growth of individual trees from a representative sample of 15 northern or southern tree species growing in a transition zone between boreal and temperate biomes. Individual trees varied in their growth response to fluctuating mean annual temperature and summer moisture stress. The variation among individuals within a species was wider than mean differences among species. The effects of mean temperature and summer moisture stress interacted, such that warm years produced positive responses to summer moisture availability and cool years produced negative responses. As climate models project significant increases in annual temperatures, growth of species like Acer saccharum, Quercus rubra, and Picea glauca will vary more in response to summer moisture stress than in the past. The magnitude of biomass growth variation in response to annual climate was 92–95% smaller than responses to tree size and age. This means that measuring or predicting the physical structure of current and future forests could tell us more about future C dynamics than growth responses related to climate change alone.     

Herbivores limit the population size of big‐leaf mahogany trees in an Amazonian forest

The Janzen–Connell hypothesis proposes that specialized herbivores maintain high numbers of tree species in tropical forests by restricting adult recruitment so that host populations remain at low densities. We tested this prediction for the large timber tree species, Swietenia macrophylla, whose seeds and seedlings are preyed upon by small mammals and a host‐specific moth caterpillar Steniscadia poliophaea, respectively. At a primary forest site, experimental seed additions to gaps – canopy‐disturbed areas that enhance seedling growth into saplings – over three years revealed lower survival and seedling recruitment closer to conspecific trees and in higher basal area neighborhoods, as well as reduced subsequent seedling survival and height growth. When we included these Janzen–Connell effects in a spatially explicit individual‐based population model, the caterpillar's impact was critical to limiting Swietenia's adult tree density, with a > 10‐fold reduction estimated at 300 years. Our research demonstrates the crucial but oft‐ignored linkage between Janzen–Connell effects on offspring and population‐level consequences for a long‐lived, potentially dominant tree species.     

The influence of sampling design on tree‐ring‐based quantification of forest growth

Tree‐rings offer one of the few possibilities to empirically quantify and reconstruct forest growth dynamics over years to millennia. Contemporaneously with the growing scientific community employing tree‐ring parameters, recent research has suggested that commonly applied sampling designs (i.e. how and which trees are selected for dendrochronological sampling) may introduce considerable biases in quantifications of forest responses to environmental change. To date, a systematic assessment of the consequences of sampling design on dendroecological and‐climatological conclusions has not yet been performed. Here, we investigate potential biases by sampling a large population of trees and replicating diverse sampling designs. This is achieved by retroactively subsetting the population and specifically testing for biases emerging for climate reconstruction, growth response to climate variability, long‐term growth trends, and quantification of forest productivity. We find that commonly applied sampling designs can impart systematic biases of varying magnitude to any type of tree‐ring‐based investigations, independent of the total number of samples considered. Quantifications of forest growth and productivity are particularly susceptible to biases, whereas growth responses to short‐term climate variability are less affected by the choice of sampling design. The world's most frequently applied sampling design, focusing on dominant trees only, can bias absolute growth rates by up to 459% and trends in excess of 200%. Our findings challenge paradigms, where a subset of samples is typically considered to be representative for the entire population. The only two sampling strategies meeting the requirements for all types of investigations are the (i) sampling of all individuals within a fixed area; and (ii) fully randomized selection of trees. This result advertises the consistent implementation of a widely applicable sampling design to simultaneously reduce uncertainties in tree‐ring‐based quantifications of forest growth and increase the comparability of datasets beyond individual studies, investigators, laboratories, and geographical boundaries.     

Strong overestimation of water‐use efficiency responses to rising CO2 in tree‐ring studies

The carbon isotope ratio (δ¹³C) in tree rings is commonly used to derive estimates of the assimilation‐to‐stomatal conductance rate of trees, that is, intrinsic water‐use efficiency (iWUE). Recent studies have observed increased iWUE in response to rising atmospheric CO₂ concentrations (C_a), in many different species, genera and biomes. However, increasing rates of iWUE vary widely from one study to another, likely because numerous covarying factors are involved. Here, we quantified changes in iWUE of two widely distributed boreal conifers using tree samples from a forest inventory network that were collected across a wide range of growing conditions (assessed using the site index, SI), developmental stages and stand histories. Using tree‐ring isotopes analysis, we assessed the magnitude of increase in iWUE after accounting for the effects of tree size, stand age, nitrogen deposition, climate and SI. We also estimated how growth conditions have modulated tree physiological responses to rising C_a. We found that increases in tree size and stand age greatly influenced iWUE. The effect of C_a on iWUE was strongly reduced after accounting for these two variables. iWUE increased in response to C_a, mostly in trees growing on fertile stands, whereas iWUE remained almost unchanged on poor sites. Our results suggest that past studies could have overestimated the CO₂ effect on iWUE, potentially leading to biased inferences about the future net carbon balance of the boreal forest. We also observed that this CO₂ effect is weakening, which could affect the future capacity of trees to resist and recover from drought episodes.     

Long Tree-Ring Chronologies Provide Evidence of Recent Tree Growth Decrease in a Central African Tropical Forest

It is still unclear whether the exponential rise of atmospheric CO₂ concentration has produced a fertilization effect on tropical forests, thus incrementing their growth rate, in the last two centuries. As many factors affect tree growth patterns, short -term studies might be influenced by the confounding effect of several interacting environmental variables on plant growth. Long-term analyses of tree growth can elucidate long-term trends of plant growth response to dominant drivers. The study of annual rings, applied to long tree-ring chronologies in tropical forest trees enables such analysis. Long-term tree-ring chronologies of three widespread African species were measured in Central Africa to analyze the growth of trees over the last two centuries. Growth trends were correlated to changes in global atmospheric CO₂ concentration and local variations in the main climatic drivers, temperature and rainfall. Our results provided no evidence for a fertilization effect of CO₂ on tree growth. On the contrary, an overall growth decline was observed for all three species in the last century, which appears to be significantly correlated to the increase in local temperature. These findings provide additional support to the global observations of a slowing down of C sequestration in the trunks of forest trees in recent decades. Data indicate that the CO₂ increase alone has not been sufficient to obtain a tree growth increase in tropical trees. The effect of other changing environmental factors, like temperature, may have overridden the fertilization effect of CO₂.     

Detecting long‐term growth trends using tree rings: a critical evaluation of methods

Tree‐ring analysis is often used to assess long‐term trends in tree growth. A variety of growth‐trend detection methods (GDMs) exist to disentangle age/size trends in growth from long‐term growth changes. However, these detrending methods strongly differ in approach, with possible implications for their output. Here, we critically evaluate the consistency, sensitivity, reliability and accuracy of four most widely used GDMs: conservative detrending (CD) applies mathematical functions to correct for decreasing ring widths with age; basal area correction (BAC) transforms diameter into basal area growth; regional curve standardization (RCS) detrends individual tree‐ring series using average age/size trends; and size class isolation (SCI) calculates growth trends within separate size classes. First, we evaluated whether these GDMs produce consistent results applied to an empirical tree‐ring data set of Melia azedarach, a tropical tree species from Thailand. Three GDMs yielded similar results – a growth decline over time – but the widely used CD method did not detect any change. Second, we assessed the sensitivity (probability of correct growth‐trend detection), reliability (100% minus probability of detecting false trends) and accuracy (whether the strength of imposed trends is correctly detected) of these GDMs, by applying them to simulated growth trajectories with different imposed trends: no trend, strong trends (?6% and +6% change per decade) and weak trends (?2%, +2%). All methods except CD, showed high sensitivity, reliability and accuracy to detect strong imposed trends. However, these were considerably lower in the weak or no‐trend scenarios. BAC showed good sensitivity and accuracy, but low reliability, indicating uncertainty of trend detection using this method. Our study reveals that the choice of GDM influences results of growth‐trend studies. We recommend applying multiple methods when analysing trends and encourage performing sensitivity and reliability analysis. Finally, we recommend SCI and RCS, as these methods showed highest reliability to detect long‐term growth trends.     

Annual wood production in a tropical rain forest in NE Costa Rica linked to climatic variation but not to increasing CO2

Increased atmospheric [CO₂] could theoretically lead to increased forest productivity (‘CO₂ fertilization’). This mechanism was hypothesized as a possible explanation for biomass increases reported from tropical forests in the last 30+ years. We used unique long‐term records of annually measured stands (eighteen 0.5 ha plots, 10 years) and focal tree species (six species, 24 years) to assess the effects of rainfall, temperature, and atmospheric [CO₂] on annual wood production in a neotropical rain forest. Our study area was a meso‐scale section (600 ha) of old‐growth Tropical Wet Forest in NE Costa Rica. Using the repeated remeasurements we directly assessed the relative effects of interannual climatic variation and increasing atmospheric [CO₂] on wood production. A remarkably simple two‐factor model explained 91% of the interannual variance in stand‐level tree growth; the statistically independent factors were total dry season rainfall (positive effect, r²=0.85) and night‐time temperature (negative effect, r²=0.42). Stand‐level tree mortality increased significantly with night‐time temperature. After accounting for dry season rainfall and night‐time temperature, there was no effect of annual [CO₂] on tree growth in either the stand or focal species data. Tree growth in this Tropical Wet Forest was surprisingly sensitive to the current range of dry season conditions and to variations in mean annual night‐time temperature of 1–2°. Our results suggest that wood production in the lowland rainforests of NE Costa Rica (and by extension in other tropical regions) may be severely reduced in future climates that are only slightly drier and/or warmer.     

Growth responses of gypsy moth larvae to elevated CO2: the influence of methods of insect rearing

Abstract The effects of elevated CO₂ on foliar chemistry of two tree species (Populus pseudo‐simonii Kitag. and Betula platyphylla) and on growth of gypsy moth (Lymantria dispar L.) larvae were examined. Furthermore, we focused on the comparison of results on the growth responses of larvae obtained from two methods of insect rearing, the no‐choice feeding trial performed in the laboratory or in situ in open‐top chambers. On the whole, both primary and secondary metabolites in the leaves of the two tree species were significantly affected by main effects of time (sampling date), CO₂ and species. Elevated CO₂ significantly increased the C : N ratio and concentrations of the soluble sugar, starch, total nonstructural carbohydrates, total phenolics and condensed tannins, but significantly decreased the concentration of nitrogen. Higher contents of total phenolics and condensed tannins were detected in the frass of larvae reared in elevated CO₂ treatments. Overall, the growth of gypsy moth larvae were significantly inhibited by elevated CO₂ and CO₂‐induced changes in leaf quality. Our study did not indicate the two methods of insect rearing could influence the direction of effects of elevated CO₂ on the growth of individual insects; however, the magnitude of negative effects of elevated CO₂ on larval growth did differ between the two insect rearing methods, and it seems that the response magnitude was also mediated by larval age and host plant species.     

Rising concentrations of atmospheric CO2 have increased growth in natural stands of quaking aspen (Populus tremuloides)

As atmospheric CO₂ levels rise, temperate and boreal forests in the Northern Hemisphere are gaining importance as carbon sinks. Quantification of that role, however, has been difficult due to the confounding effects of climate change. Recent large‐scale experiments with quaking aspen (Populus tremuloides), a dominant species in many northern forest ecosystems, indicate that elevated CO₂ levels can enhance net primary production. Field studies also reveal that droughts contribute to extensive aspen mortality. To complement this work, we analyzed how the growth of wild aspen clones in Wisconsin has responded to historical shifts in CO₂ and climate, accounting for age, genotype (microsatellite heterozygosity), and other factors. Aspen growth has increased an average of 53% over the past five decades, primarily in response to the 19.2% rise in ambient CO₂ levels. CO₂‐induced growth is particularly enhanced during periods of high moisture availability. The analysis accounts for the highly nonlinear changes in growth rate with age, and is unaffected by sex or location sampled. Growth also increases with individual heterozygosity, but this heterozygote advantage has not changed with rising levels of CO₂ or moisture. Thus, increases in future growth predicted from previous large‐scale, common‐garden work are already evident in this abundant and ecologically important tree species. Owing to aspen's role as a foundation species in many North American forest ecosystems, CO₂‐stimulated growth is likely to have repercussions for numerous associated species and ecosystem processes.     

Long‐term physiological and growth responses of Himalayan fir to environmental change are mediated by mean climate

High‐elevation forests are experiencing high rates of warming, in combination with CO₂ rise and (sometimes) drying trends. In these montane systems, the effects of environmental changes on tree growth are also modified by elevation itself, thus complicating our ability to predict effects of future climate change. Tree‐ring analysis along an elevation gradient allows quantifying effects of gradual and annual environmental changes. Here, we study long‐term physiological (ratio of internal to ambient CO₂, i.e., C_i/C_a and intrinsic water‐use efficiency, iWUE) and growth responses (tree‐ring width) of Himalayan fir (Abies spectabilis) trees in response to warming, drying, and CO₂ rise. Our study was conducted along elevational gradients in a dry and a wet region in the central Himalaya. We combined dendrochronology and stable carbon isotopes (δ¹³C) to quantify long‐term trends in C_i/C_a ratio and iWUE (δ¹³C‐derived), growth (mixed‐effects models), and evaluate climate sensitivity (correlations). We found that iWUE increased over time at all elevations, with stronger increase in the dry region. Climate–growth relations showed growth‐limiting effects of spring moisture (dry region) and summer temperature (wet region), and negative effects of temperature (dry region). We found negative growth trends at lower elevations (dry and wet regions), suggesting that continental‐scale warming and regional drying reduced tree growth. This interpretation is supported by δ¹³C‐derived long‐term physiological responses, which are consistent with responses to reduced moisture and increased vapor pressure deficit. At high elevations (wet region), we found positive growth trends, suggesting that warming has favored tree growth in regions where temperature most strongly limits growth. At lower elevations (dry and wet regions), the positive effects of CO₂ rise did not mitigate the negative effects of warming and drying on tree growth. Our results raise concerns on the productivity of Himalayan fir forests at low and middle (<3,300 m) elevations as climate change progresses.     

Differential effects of elevated CO2 on acorn density, weight, germination, and predation among three oak species in a scrub-oak forest

Much research on the effects of elevated CO₂ on forest trees has focused on quantitative changes in photosynthesis, secondary chemistry, and plant biomass. However, plant fitness responses to rising CO₂ should also include quantitative measures of reproduction, since most forest systems are recruitment limited. Until now, it has proved very difficult to grow forest trees to sexual maturity in a CO₂‐enriched environment. This paper is the first of its kind to address the effects of elevated CO₂ on the reproduction of hardwood trees in a natural forest. Beginning in 1996, scrub‐oak vegetation, predominantly three species of scrub‐oaks, Quercus myrtifolia, Q. chapmanii, and Q. geminata, were grown inside eight chambers with elevated CO₂ (704 parts per million (ppm)) and eight with ambient CO₂ (379 ppm) at Kennedy Space Center, Florida. In elevated CO₂, acorn production increased significantly for the dominant species Q. myrtifolia and for Q. chapmanii, but it did not increase for the subdominant, Q. geminata. Acorn weight, germination rate, and predation by weevils were unaffected by CO₂. Thus, recruitment of some forest tree species into the Florida scrub‐oak community is likely to be accelerated in an atmosphere of increased CO₂. However, because the acceleration of recruitment differs among species, over the long term, Q. myrtifolia and Q. chapmanii will be favored over Q. geminata and this is likely to change patterns of species diversity.     

Chronic water stress reduces tree growth and the carbon sink of deciduous hardwood forests

Predicted decreases in water availability across the temperate forest biome have the potential to offset gains in carbon (C) uptake from phenology trends, rising atmospheric CO₂, and nitrogen deposition. While it is well established that severe droughts reduce the C sink of forests by inducing tree mortality, the impacts of mild but chronic water stress on forest phenology and physiology are largely unknown. We quantified the C consequences of chronic water stress using a 13‐year record of tree growth (n = 200 trees), soil moisture, and ecosystem C balance at the Morgan–Monroe State Forest (MMSF) in Indiana, and a regional 11‐year record of tree growth (n > 300 000 trees) and water availability for the 20 most dominant deciduous broadleaf tree species across the eastern and midwestern USA. We show that despite ~26 more days of C assimilation by trees at the MMSF, increasing water stress decreased the number of days of wood production by ~42 days over the same period, reducing the annual accrual of C in woody biomass by 41%. Across the deciduous forest region, water stress induced similar declines in tree growth, particularly for water‐demanding ‘mesophytic’ tree species. Given the current replacement of water‐stress adapted ‘xerophytic’ tree species by mesophytic tree species, we estimate that chronic water stress has the potential to decrease the C sink of deciduous forests by up to 17% (0.04 Pg C yr^?1) in the coming decades. This reduction in the C sink due to mesophication and chronic water stress is equivalent to an additional 1–3 days of global C emissions from fossil fuel burning each year. Collectively, our results indicate that regional declines in water availability may offset the growth‐enhancing effects of other global changes and reduce the extent to which forests ameliorate climate warming.     

Growth, carbon-isotope discrimination, and drought-associated mortality across a Pinus ponderosa elevational transect

Drought‐ and insect‐associated tree mortality at low‐elevation ecotones is a widespread phenomenon but the underlying mechanisms are uncertain. Enhanced growth sensitivity to climate is widely observed among trees that die, indicating that a predisposing physiological mechanism(s) underlies tree mortality. We tested three, linked hypotheses regarding mortality using a ponderosa pine (Pinus ponderosa) elevation transect that experienced low‐elevation mortality following prolonged drought. The hypotheses were: (1) mortality was associated with greater growth sensitivity to climate, (2) mortality was associated with greater sensitivity of gas exchange to climate, and (3) growth and gas exchange were correlated. Support for all three hypotheses would indicate that mortality results at least in part from gas exchange constraints. We assessed growth using basal area increment normalized by tree basal area [basal area increment (BAI)/basal area (BA)] to account for differences in tree size. Whole‐crown gas exchange was indexed via estimates of the CO₂ partial pressure difference between leaf and atmosphere (p_a?p_c) derived from tree ring carbon isotope ratios (δ¹³C), corrected for temporal trends in atmospheric CO₂ and δ¹³C and elevation trends in pressure. Trees that survived the drought exhibited strong correlations among and between BAI, BAI/BA, p_a?p_c, and climate. In contrast, trees that died exhibited greater growth sensitivity to climate than trees that survived, no sensitivity of p_a?p_c to climate, and a steep relationship between p_a?p_c and BAI/BA. The p_a?p_c results are consistent with predictions from a theoretical hydraulic model, suggesting trees that died had a limited buffer between mean water availability during their lifespan and water availability during drought – i.e., chronic water stress. It appears that chronic water stress predisposed low‐elevation trees to mortality during drought via constrained gas exchange. Continued intensification of drought in mid‐latitude regions may drive increased mortality and ecotone shifts in temperate forests and woodlands.