Leaf-out Phenology Research Articles

Elevated CO2 does not improve seedling performance in a naturally regenerated oak woodland exposed to biotic stressors

IntroductionInsect herbivores and biotrophic pathogens are major stressors influencing natural regeneration in woodlands. Information on the effect of elevated CO2 (eCO2) on plant-insect-pathogen interactions under natural conditions is lacking.MethodsWe studied the effects of eCO2 on leaf-out phenology, as well as on levels of insect herbivory and powdery mildew (PM), i.e., reduction of leaf photosynthetic material. We then assessed the combined impacts of these biotic stressors and eCO2 on seedling photosynthesis and growth. A total of 92 naturally recruited and 114 potted seedlings of 5 temperate tree species (Quercus robur, Acer pseudoplatanus, Corylus avellana, Crataegus monogyna, and Ilex aquifolium) within a mature oak woodland were studied.ResultsWe found that eCO2 advanced leaf-out phenology and was a significant explanatory variable for growth and physiological performance in potted seedlings. Potted oak seedlings experienced 11-fold higher tissue loss from insect herbivory than natural seedlings. The earliest leaf-flushing species, hawthorn, and the evergreen holly were resistant to insect attack and were not affected by PM. Oak was defoliated most but showed the highest regeneration capacity. Hazel was more resistant to PM infection than oak and sycamore. Despite being highly infected by PM, sycamore was less affected than oak. The more vigorous sycamore and oak seedlings suffered more severe PM disease.ConclusionNo evidence emerged that eCO2 enhances natural regeneration under biotic stress for any of the species studied.

Genetic Differentiation of Budburst Timing in Fagus crenata Populations along a Spatial Gradient in Late Frost Timing in the Hakkoda Mountains, Northern Japan

We studied the genetic differentiation in budburst timing among Fagus crenata populations along spatial gradients in late frost timing in the Hakkoda Mountains, northern Japan, by focusing on last fatal frost day and topography. For budburst timing, we analyzed interpopulation variations in habitats, genetic variations in a nursery, and the relationships between these variations and environmental conditions in the habitats. Analyses of interpopulation variation showed that the day and the temperature sum of budburst positively correlated with the last fatal frost day in the habitats. Analyses of genetic variation showed significant genetic variations among provenances and families for both traits. For all provenances, the heritability for these traits were 0.7–0.8. The genetic variations were significantly associated with variations in the last fatal frost day among the provenances, suggesting that natural selection due to late fatal frost causes genetic differentiation in the traits along the spatial gradient in late frost timing. These results demonstrate that late frost is a key factor driving genetic differentiation of leaf-out phenology within a regional tree population.

Stacked distribution models predict climate-driven loss of variation in leaf phenology at continental scales

Climate change is having profound effects on species distributions and is likely altering the distribution of genetic variation across landscapes. Maintaining population genetic diversity is essential for the survival of species facing rapid environmental change, and variation loss will further ecological and evolutionary change. We used trait values of spring foliar leaf-out phenology of 400 genotypes from three geographically isolated populations of Populus angustifolia grown under common conditions, in concert with stacked species distribution modeling, to ask: (a) How will climate change alter phenological variation across the P. angustifolia species-range, and within populations; and (b) will the distribution of phenological variation among and within populations converge (become more similar) in future climatic conditions? Models predicted a net loss of phenological variation in future climate scenarios on 20-25% of the landscape across the species’ range, with the trailing edge population losing variation on as much as 47% of the landscape. Our models also predicted that population’s phenological trait distributions will become more similar over time. This stacked distribution model approach allows for the identification of areas expected to experience the greatest loss of genetically based functional trait variation and areas that may be priorities to conserve as future genetic climate refugia.

Differences between four sympatric subtropical tree species in the interactive effects of three environmental cues on leaf-out phenology

Climatic warming is currently changing the spring phenology of extratropical trees, and this has several important effects on the trees and ecosystems. The major climatic cues regulating the spring phenology are winter chilling, spring forcing, and photoperiod. The interactions between these three remain largely unstudied because most studies concentrate on the effects of one cue, or maximally two, at a time. We studied the effects and interactions of chilling duration, forcing temperature, and forcing photoperiod simultaneously in four subtropical tree species. The main emphasis in our experiments was on the interaction of chilling duration and forcing temperature. The existence of this interaction was suggested in the ‘Vegis theory’, put forward decades ago but largely forgotten since. We also introduced a novel method for testing the theory experimentally. We found support for the Vegis theory in two of the four species examined. In the other two species the leaf-out timing was largely controlled by spring forcing. The effects of photoperiod were generally minor. Our results show that there are major differences between sympatric subtropical tree species in their phenological responses to environmental cues. These differences need to be addressed in the development of process-based tree phenology models. Our results further suggest that different subtropical trees respond differently to climatic warming because of differences related to the Vegis theory. This hypothesis remains to be tested in further studies.

Winter warming offsets one half of the spring warming effects on leaf unfolding.

Winter temperature‐related chilling and spring temperature‐related forcing are two major environmental cues shaping the leaf‐out date of temperate species. To what degree insufficient chilling caused by winter warming would slow phenological responses to spring warming remains unclear. Using 27,071 time series of leaf‐out dates for 16 tree species in Europe, we constructed a phenological model based on the linear or exponential function between the chilling accumulation (CA) and forcing requirements (FR) of leaf‐out. We further used the phenological model to quantify the relative contributions of chilling and forcing on past and future spring phenological change. The results showed that the delaying effect of decreased chilling on the leaf‐out date was prevalent in natural conditions, as more than 99% of time series exhibited a negative relationship between CA and FR. The reduction in chilling linked to winter warming from 1951 to 2014 could offset about one half of the spring phenological advance caused by the increase in forcing. In future warming scenarios, if the same model is used and a linear, stable correlation between CA and FR is assumed, declining chilling will continuously offset the advance of leaf‐out to a similar degree. Our study stresses the importance of assessing the antagonistic effects of winter and spring warming on leaf‐out phenology.

Climate Warming Increased Spring Leaf-Out Variation Across Temperate Trees in China

Leaf-out phenology plays a key role in ecosystem structure and functioning. Phenological changes have often been linked to climatic factors and have received considerable attention, with most studies focusing on trends of leaf-out phenology. Leaf-out variation (LOV), which reflects the stability of phenological responses, may also be affected by climate change, yet this has received less scientific attention. In this study, we examined spring LOV in response to climate change in China during the period 1963–2008 using in situ records of 15 species at 25 phenological observation sites across several climate zones and explored spatiotemporal changes of LOV and the underlying mechanisms. We observed a significant decrease of LOV toward higher latitudes (−0.2 ± 0.1 days⋅°N–1;P < 0.001) across all species. Temporally, we found that the LOV was significantly increased from the period 1963–1986 (6.9 ± 2.8 days) to the period 1987–2008 (7.9 ± 3.7 days, P < 0.05). Furthermore, the LOV changes between 1987–2008 and 1963–1986 were significantly smaller at high latitudes (average decrease of 1.0 day) than at low latitudes (average increase of 1.5 days). The spatial pattern of LOV is likely due to both increased heat requirements and greater temperature sensitivity at low latitudes compared with high latitudes. The temporal pattern of LOV is likely related to increased heat requirements for leaf-out during 1987–2008 when the average air temperature was higher. Our analysis indicated that the phenology response to climate change is reflected not only in the temporal trends for long time series but also in the variation of phenological dates. Results from this study improve our understanding of phenological responses to climate change and could be applied in the assessment of regional phenology changes to evaluate better the impacts of climate change on ecosystem structure and function.

Extending the Cultivation Area of Pecan (Carya illinoinensis) Toward the South in Southeastern Subtropical China May Cause Increased Cold Damage.

Pecan (Carya illinoinensis) is an important nut tree species in its native areas in temperate and subtropical North America, and as an introduced crop in subtropical southeastern China as well. We used process-based modeling to assess the effects of climatic warming in southeastern China on the leaf-out phenology of pecan seedlings and the subsequent risk of “false springs,” i.e., damage caused by low temperatures occurring as a result of prematurely leafing out. In order to maximize the biological realism of the model used in scenario simulations, we developed the model on the basis of experiments explicitly designed for determining the responses modeled. The model showed reasonable internal accuracy when calibrated against leaf-out observations in a whole-tree chamber (WTC) experiment with nine different natural-like fluctuating temperature treatments. The model was used to project the timing of leaf-out in the period 2022–2099 under the warming scenarios RCP4.5 and RCP8.5 in southeastern China. Two locations in the main pecan cultivation area in the northern subtropical zone and one location south of the main cultivation area were addressed. Generally, an advancing trend of leaf-out was projected for all the three locations under both warming scenarios, but in the southern location, a delay was projected under RCP8.5 in many years during the first decades of the 21st century. In the two northern locations, cold damage caused by false springs was projected to occur once in 15–26 years at most, suggesting that pecan cultivation can be continued relatively safely in these two locations. Paradoxically, more frequent cold damage was projected for the southern location than for the two northern locations. The results for the southern location also differed from those for the northern locations in that more frequent cold damage was projected under the RCP4.5 warming scenario (once in 6 years) than under the RCP8.5 scenario (once in 11 years) in the southern location. Due to the uncertainties of the model applied, our conclusions need to be re-examined in an additional experimental study and further model development based on it; but on the basis of our present results, we do not recommend starting large-scale pecan cultivation in locations south of the present main pecan cultivation area in southeastern subtropical China.

Improved phenological escape can help temperate tree seedlings maintain demographic performance under climate change conditions.

Phenological escape, a strategy that deciduous understory plants use to access direct light in spring by leafing out before the canopy closes, plays an important role in shaping the recruitment of temperate tree seedlings. Previous studies have investigated how climate change will alter these dynamics for herbaceous species, but there is a knowledge gap related to how woody species such as tree seedlings will be affected. Here, we modeled temperate tree seedling leaf-out phenology and canopy close phenology in response to environmental drivers and used climate change projections to forecast changes to the duration of spring phenological escape. We then used these predictions to estimate changes in annual carbon assimilation while accounting for reduced carbon assimilation rates associated with hotter and drier summers. Lastly, we applied these estimates to previously published models of seedling growth and survival to investigate the net effect on seedling demographic performance. Our models predict that temperate tree seedlings will experience improved phenological escape and, therefore, increased spring carbon assimilation under climate change conditions. However, increased summer respiration costs will offset the gains in spring under extreme climate change leading to a net loss in annual carbon assimilation and demographic performance. Furthermore, we found that annual carbon assimilation predictions depend strongly on the species of nearby canopy tree that seedlings were planted near, with all seedlings projected to assimilate less carbon (and therefore experience worse demographic performance) when planted near Quercus rubra canopy trees as opposed to Acer saccharum canopy trees. We conclude that changes to spring phenological escape will have important effects on how tree seedling recruitment is affected by climate change, with the magnitude of these effects dependent upon climate change severity and biological interactions with neighboring adults. Thus, future studies of temperate forest recruitment should account for phenological escape dynamics in their models.

Phylogenetic conservatism in heat requirement of leaf-out phenology, rather than temperature sensitivity, in Tibetan Plateau

Phylogenetic studies on the leaf unfolding date are expected to improve understanding of phenological change. However, it remains unclear whether the response of leaf unfolding date to temperature is phylogenetically conserved among close relatives across differential and harsh habitats. Meanwhile, the phenological mechanism underlying the role of phylogenetic relationship on plant phenology remains unclear. We used Blomberg's K to assess phylogenetic signal in leaf unfolding date, its sensitivities to daily mean, minimum and maximum temperatures, in growing-degree-days and average level of each of daily mean, minimum and maximum temperatures experienced by plants over a given period prior to leaf unfolding date for eighteen species at seventeen sites in the Tibetan Plateau. We showed that leaf unfolding date and its sensitivities to daily mean, minimum and maximum temperatures did not exhibit overall significant phylogenetic signals. Moreover, the sensitivity of leaf unfolding date to daily minimum temperature exhibited phylogenetic antisignal at species level, which indicated the trait difference among close relatives exceed that among distantly related lineages. However, there were overall significant phylogenetic signals in growing-degree-days and the average level of each of daily mean, minimum and maximum temperatures experienced by plants before leaf unfolding date. The signals were stronger for the growing-degree-days and the average level of each of daily mean, minimum and maximum temperatures in the period closer to leaf unfolding date. Our results imply that close relatives trace similar heat requirements for leaf unfolding date in the harsh and spatially heterogeneous environment of the Tibetan Plateau. However, the time when these lineage-specific heat requirements are met varies spatially depending on local conditions, thereby resulting in phylogenetic non-conservatism of leaf unfolding date and its temperature sensitivities. This provides a phenological explanation, for the first time, for phylogenetic non-conservatism in plant leafing phenology.

Spatiotemporal variations in leaf-out phenology of typical European tree species and their responses to climate change.

Over the past decades, global warming significantly affected the spring phenology of plants. Many studies have reported the temporal and spatial patterns of spring phenological changes in China, but relatively less is known for that in Europe, which is also located in the temperate area of the Northern Hemisphere. To facilitate the regional comparison of phenological change and understand its response to climate change, we used the data of first leaf date (FLD) in Europe (1980-2014) and the corresponding meteorological data to examine the spatiotemporal variations in leaf-out phenology of four typical tree species (Aesculus hippocastanum, Betula pendula, Fagus sylvatica, and Quercus robur), and to identify the major climatic factors driving such variations. The results showed that the FLD of the four species in the study area advanced by 3.3-7.5 d·10 a-1 during 1980-2014. The FLD was delayed at a rate of 2.03-3.19 d per degree of latitude from south to north, of 0.19-0.80 d per degree of longitude from west to East (except for Fagus sylvatica), of 2.25-3.44 d·100 m-1 from low to high elevation. The advances in the FLD were mainly attributed to the increases of temperature in spring and the increases of precipitation in spring and winter. The rise of temperature in autumn and winter would delay FLD.

Divergent carbon cycle response of forest and grass-dominated northern temperate ecosystems to record winter warming.

Northern temperate ecosystems are experiencing warmer and more variable winters, trends that are expected to continue into the foreseeable future. Despite this, most studies have focused on climate change impacts during the growing season, particularly when comparing responses across different vegetation cover types. Here we examined how a perennial grassland and adjacent mixed forest ecosystem in New Hampshire, United States, responded to a period of highly variable winters from 2014 through 2017 that included the warmest winter on record to date. In the grassland, record-breaking temperatures in the winter of 2015/2016 led to a February onset of plant growth and the ecosystem became a sustained carbon sink well before winter ended, taking up roughly 90g/m2 more carbon during the winter to spring transition than in other recorded years. The forest was an unusually large carbon source during the same period. While forest photosynthesis was restricted by leaf-out phenology, warm winter temperatures caused large pulses of ecosystem respiration that released nearly 230gC/m2 from February through April, more than double the carbon losses during that period in cooler years. These findings suggest that, as winters continue to warm, increases in ecosystem respiration outside the growing season could outpace increases in carbon uptake during a longer growing season, particularly in forests that depend on leaf-out timing to initiate carbon uptake. In ecosystems with a perennial leaf habit, warming winter temperatures are more likely to increase ecosystem carbon uptake through extension of the active growing season. Our results highlight the importance of understanding relationships among antecedent winter conditions and carbon exchange across land-cover types to understand how landscape carbon exchange will change under projected climate warming.

An introduced plant is associated with declines in terrestrial arthropods, but no change in stream invertebrates

Riverine systems often spread non-native species, yet the co-occurring impacts of introduced riparian vegetation on aquatic- and terrestrial-derived resources are unknown. We compared aquatic and terrestrial arthropod communities and their flux into and out of streams in riparian reaches invaded and uninvaded by Robinia neomexicana, a woody plant introduced to a western Colorado watershed. We found that invaded reaches had fewer terrestrial arthropods collected off foliage, conceivably because of the plant’s later leaf-out phenology. Overall, seasonal and annual factors best described terrestrial and aquatic arthropod communities. However, when we evaluated vegetation and stream characteristics in lieu of season and year, we found terrestrial arthropod biomass and richness were negatively related to cover of R. neomexicana and positively related to vegetative cover, forb cover, and vertical vegetation structure. Our results suggest ecosystems respond to landscape variation differently, where directly related food web components (i.e., terrestrial arthropods on introduced vegetation) respond stronger than more distally related constituents (i.e., aquatic insects).

Daylength helps temperate deciduous trees to leaf-out at the optimal time.

Global warming has led to substantially earlier spring leaf-out in temperate-zone deciduous trees. The interactive effects of temperature and daylength underlying this warming response remain unclear. However, they need to be accurately represented by earth system models to improve projections of the carbon and energy balances of temperate forests and the associated feedbacks to the Earth's climate system. We studied the control of leaf-out by daylength and temperature using data from six tree species across 2,377 European phenological network (www.pep725.eu), each with at least 30years of observations. We found that, in addition to and independent of the known effect of chilling, daylength correlates negatively with the heat requirement for leaf-out in all studied species. In warm springs when leaf-out is early, days are short and the heat requirement is higher than in an average spring, which mitigates the warming-induced advancement of leaf-out and protects the tree against precocious leaf-out and the associated risks of late frosts. In contrast, longer-than-average daylength (in cold springs when leaf-out is late) reduces the heat requirement for leaf-out, ensuring that trees do not leaf-out too late and miss out on large amounts of solar energy. These results provide the first large-scale empirical evidence of a widespread daylength effect on the temperature sensitivity of leaf-out phenology in temperate deciduous trees.

Local environment, not local adaptation, drives leaf-out phenology in common gardens along an elevational gradient in Acadia National Park, Maine.

Climate-driven changes in phenology are substantially affecting ecological relationships and ecosystem processes. The role of variation among species has received particular attention; for example, variation among species' phenological responses to climate can disrupt trophic interactions and can influence plant performance. Variation within species in phenological responses to climate, however, has received much less attention, despite its potential role in ecological interactions and local adaptation to climate change. We constructed three common gardens across an elevation gradient on Cadillac Mountain in Acadia National Park, Maine, to test population-level responses in leaf-out phenology in a reciprocal transplant experiment. The experiment included three native species: low bush blueberry (Vaccinium angustifolium), sheep's laurel (Kalmia angustifolia), and three-toothed cinquefoil (Sibbaldiopsis tridentata). Evidence for local adaptation of phenological response to temperature varied among the species, but was weak for all three. Rather, variation in phenological response to temperature appeared to be driven by local microclimate at each garden site and year-to-year variation in temperature. Population-level adaptations in leaf-out phenology appear to be relatively unimportant for these species in Acadia National Park, perhaps a reflection of strong genetic mixing across elevations, or weak differences in selection on phenological response to spring temperatures at different elevations. These results concur with other observational data in Acadia and highlight the utility of experimental approaches to understand the importance of annual and local site variation in affecting phenology both among and within plant species.

From observations to experiments in phenology research: investigating climate change impacts on trees and shrubs using dormant twigs.

Climate change is advancing the leaf-out times of many plant species and mostly extending the growing season in temperate ecosystems. Laboratory experiments using twig cuttings from woody plant species present an affordable, easily replicated approach to investigate the relative importance of factors such as winter chilling, photoperiod, spring warming and frost tolerance on the leafing-out times of plant communities. This Viewpoint article demonstrates how the results of these experiments deepen our understanding beyond what is possible via analyses of remote sensing and field observation data, and can be used to improve climate change forecasts of shifts in phenology, ecosystem processes and ecological interactions. The twig method involves cutting dormant twigs from trees, shrubs and vines on a single date or at intervals over the course of the winter and early spring, placing them in containers of water in controlled environments, and regularly recording leaf-out, flowering or other phenomena. Prior to or following leaf-out or flowering, twigs may be assigned to treatment groups for experiments involving temperature, photoperiod, frost, humidity and more. Recent studies using these methods have shown that winter chilling requirements and spring warming strongly affect leaf-out and flowering times of temperate trees and shrubs, whereas photoperiod requirements are less important than previously thought for most species. Invasive plant species have weaker winter chilling requirements than native species in temperate ecosystems, and species that leaf-out early in the season have greater frost tolerance than later leafing species. This methodology could be extended to investigate additional drivers of leaf-out phenology, leaf senescence in the autumn, and other phenomena, and could be a useful tool for education and outreach. Additional ecosystems, such as boreal, southern hemisphere and sub-tropical forests, could also be investigated using dormant twigs to determine the drivers of leaf-out times and how these ecosystems will be affected by climate change.

Consequences of phenology variation and oxidative defenses in Quercus

Ecologists are interested in consequences of changes in the timing of ecological interactions because of the potential for widespread phenological shifts under climate change. Plant–herbivore interactions may be particularly affected by phenology changes if they result in differences in leaf defensive chemistry and other aspects of host foliage quality. Recent studies of defensive leaf phenolics have focused on their ability to create oxidative stress in herbivores, but the phenology of this “oxidative capacity,” including seasonal patterns and how it is affected by leaf-out phenology, is unknown. We combined observational and experimental studies of oaks (Quercus spp.) to determine if leaf-out phenology correlates with oxidative capacity and leaf characteristics, herbivore abundance, and leaf damage, and how oxidative capacity changes through spring and summer leaf development. Oaks with earlier leaf-out had significantly lower oxidative capacity in spring and early summer, and there was a trend for higher spring leaf damage on these trees. One group of spring herbivores, leaf rollers, was more abundant on earlier-leafing trees. By mid-summer, leaf quality differences disappeared. Repeated measurements of individual trees demonstrated that oxidative capacity is highest shortly after leaf-out but declines within 2 weeks. These results partially contrast traditional views of leaf defense phenology where leaves are most susceptible shortly after leaf-out, but defenses build as proanthocyanidins accumulate in expanded leaves. Early season oxidative defenses may have a significant effect on plant growth by protecting young leaves at a time when leaf tissue loss will have the greatest consequences for later photosynthesis.

Drivers of leaf‐out phenology and their implications for species invasions: insights from Thoreau's Concord

To elucidate climate-driven changes in leaf-out phenology and their implications for species invasions, we observed and experimentally manipulated leaf out of invasive and native woody plants in Concord, MA, USA. Using observations collected by Henry David Thoreau (1852-1860) and our own observations (2009-2013), we analyzed changes in leaf-out timing and sensitivity to temperature for 43 woody plant species. We experimentally tested winter chilling requirements of 50 species by exposing cut branches to warm indoor temperatures (22°C) during the winter and spring of 2013. Woody species are now leafing out an average of 18d earlier than they did in the 1850s, and are advancing at a rate of 5±1d°C(-1) . Functional groups differ significantly in the duration of chilling they require to leaf out: invasive shrubs generally have weaker chilling requirements than native shrubs and leaf out faster in the laboratory and earlier in the field; native trees have the strongest chilling requirements. Our results suggest that invasive shrub species will continue to have a competitive advantage as the climate warms, because native plants are slower to respond to warming spring temperatures and, in the future, may not meet their chilling requirements.

Mapping asynchrony between gypsy moth egg-hatch and forest leaf-out: Putting the phenological window hypothesis in a spatial context

Spring phenology in temperate forest ecosystems is responding in diverse ways to global climate change, with unknown consequences for insect disturbances that affect forest productivity. Adaptive forest management that anticipates changes in insect disturbance regimes requires an understanding of the mechanistic links between climate and disturbance behavior. Phenological mismatches between insects and their hosts have the potential to modify patterns of defoliation that are a significant disturbance in temperate forests. Yet the frequency and extent of phenological mismatches have not been quantified over forest landscapes. Forest dwelling, spring-feeding defoliators must synchronize larval emergence annually with leaf-out phenology of host trees. Lack of synchrony can limit population expansion, unless dispersing larvae successfully locate phenologically compatible hosts. We quantified the potential for successful dispersal of an introduced defoliator in a forest landscape by mapping phenological asynchrony from 2000 to 2010. Maps compared MODIS derived leaf-out with gypsy moth egg-hatch from the BioSIM insect phenology model. Mean temporal differences between gypsy moth egg-hatch and leaf-out were sufficient (<−5 or >+10days) to severely limit 1st instar survival in 5 of 11years. Biologically meaningful phenological asynchrony extended over distances of 3.1–4.9km, well beyond typical gypsy moth dispersal thresholds (<1km), meaning that larvae in some parts of the landscape were unable to escape unfavorable phenology by dispersing. Significant spatial heterogeneity in phenological asynchrony clarifies its potential to affect gypsy moth population dynamics and patterns of defoliation disturbance, in contrast with previous field-based research conducted at smaller spatial scales. Our results suggest that delay of leaf-out in this ecosystem due to inadequate winter chilling may disrupt gypsy moth disturbances, as years where host forest phenology is significantly asynchronous over a majority of the landscape become more common. Our modeling approach is relevant and applicable to a wide range of ecological phenomena that require synchrony of phenological interactions across trophic levels. The results inform forest management by illustrating areas in host dominated landscapes that are most susceptible to defoliation in a given year, allowing for improved planning of insect suppression activities.

Predicting effects of global warming on growth and mortality of upland oak species in the midwestern United States: a physiologically based dendroecological approach

This study combined an ecophysiological model and dendroecological analyses to evaluate potential effects of global warming on the physiology, growth, and mortality of white oak (Quercusalba L.) and black oak (Quercusvelutina Lam.) in the Ohio River region. The model integrated data for ecophysiology of oak species, site attributes, and daily temperature and precipitation to model nonlinear responses of stomatal conductance (gs), net photosynthesis (Pnet), and woody respiration (Rw) to variations in temperature and soil water content. Relationships between modeled physiological response indices and actual white and black oak annual radial growth indices were evaluated by regression analyses, using growth and weather data for the period 1900–1987 for seven upland oak–hickory forests. Modeled physiological response indices explained 40–60% of variation in radial growth indices. To evaluate the effects of global warming, daily temperature values for the period 1900–1987 were increased by 2 or 5 °C, without changing precipitation values, and physiological response indices were computed. Model indices generated in warming simulations were entered into dendroclimatic regression models calibrated under conditions without any warming to predict radial growth under warming scenarios. Under the warming scenarios, OAKWBAL predicted a substantial increase in growing season Rw, but little change in growing season Pnet. Warming merely shifted the period of near-maximal Pnet earlier in the growing season, without changing its duration. However, this result was somewhat dependent upon the ability of leaf-out phenology to track changes in temperature regime. The net effect of increased Rw, with little change in Pnet, was a reduction in radial growth and a higher frequency of years with climatic conditions stressful to oaks on upland sites. A historical association between severe drought and increased incidence of oak growth decline and mortality indicated that global warming could increase the incidence of decline and mortality in oak populations on upland sites similar to those in this study.