Predicting phenology by integrating ecology,evolution and climate science

Forecasting how species and ecosystems will respond to climate change has been a major aim of ecology in recent years. Much of this research has focused on phenology – the timing of life‐history events. Phenology has well‐demonstrated links to climate, from genetic to landscape scales; yet our ability to explain and predict variation in phenology across species, habitats and time remains poor. Here, we outline how merging approaches from ecology, climate science and evolutionary biology can advance research on phenological responses to climate variability. Using insight into seasonal and interannual climate variability combined with niche theory and community phylogenetics, we develop a predictive approach for species’ reponses to changing climate. Our approach predicts that species occupying higher latitudes or the early growing season should be most sensitive to climate and have the most phylogenetically conserved phenologies. We further predict that temperate species will respond to climate change by shifting in time, while tropical species will respond by shifting space, or by evolving. Although we focus here on plant phenology, our approach is broadly applicable to ecological research of plant responses to climate variability.     

A parameterization of leaf phenology for the terrestrial ecosystem component of climate models

Leaf phenology remains one of the most difficult processes to parameterize in terrestrial ecosystem models because our understanding of the physical processes that initiate leaf onset and senescence is incomplete. While progress has been made at the molecular level, for example by identifying genes that are associated with senescence and flowering for selected plant species, a picture of the processes controlling leaf phenology is only beginning to emerge. A variety of empirical formulations have been used with varying degrees of success in terrestrial ecosystem models for both extratropical and tropical biomes. For instance, the use of growing degree‐days (GDDs) to initiate leaf onset has received considerable recognition and this approach is used in a number of models. There are, however, limitations when using GDDs and other empirically based formulations in global transient climate change simulations. The phenology scheme developed for the Canadian Terrestrial Ecosystem Model (CTEM), designed for inclusion in the Canadian Centre for Climate Modelling and Analysis coupled general circulation model, is described. The representation of leaf phenology is general enough to be applied over the globe and sufficiently robust for use in transient climate change simulations. Leaf phenology is functionally related to the (possibly changing) climate state and to atmospheric composition rather than to geographical boundaries or controls implicitly based on current climate. In this approach, phenology is controlled by environmental conditions as they affect the carbon balance. A carbon‐gain‐based scheme initiates leaf onset when it is beneficial for the plant, in carbon terms, to produce new leaves. Leaf offset is initiated by unfavourable environmental conditions that incur carbon losses and these include shorter day length, cooler temperatures, and dry soil moisture conditions. The comparison of simulated leaf onset and offset times with observation‐based estimates for temperate and boreal deciduous, tropical evergreen, and tropical deciduous plant functional types at selected locations indicates that the phenology scheme performs satisfactorily. Model simulated leaf area index and stem and root biomass are also compared with observational estimates to illustrate the performance of CTEM.     

Exploring the impact of neutral evolution on intrapopulation genetic differentiation in functional traits in a long-lived plant

Most plant species, particularly long-lived plants, harbor a large amount of genetic variation within populations. A central issue in evolutionary ecology is to explore levels of genetic variation and understand the mechanisms that influence them. In this study, our goals were to examine the impact of neutral evolutionary processes on the genetic variance and functional diversity within three populations of a long-lived plant (Quercus suber L.). For this purpose, we genotyped the progeny of 45 open-pollinated mother trees from three populations originating from Spain, Portugal, and Morocco using six microsatellite markers. Seedlings were planted in a common garden trial and were phenotypically characterized by seven leaf functional traits. Molecular analyses revealed weak genetic differences between Iberian and Moroccan populations. Nevertheless, high genetic differentiation was observed among maternal families within populations. Differentiation between particular maternal families from the same population reached values of 29.2 %, which far exceeds the values reported between the most genetically distant populations for this species (11.7 %). Maternal families differed also in phenology, leaf size, and shape traits. In the Moroccan population, there were correlations among matrices of distances for molecular markers, leaf shape traits (e.g., leaf circularity index), and phenology, indicating that maternal families with contrasting phenologies were genetically and functionally distinct. This, together with the moderate heritability for phenology in Moroccan population, suggests that besides selective forces, neutral evolutionary processes have promoted intrapopulation genetic divergence and contribute to maintain high levels of genetic variation within this population. Overall, our results reinforce the importance of intrapopulation studies in long-lived plants under an evolutionary context.     

Macrophenology: insights into the broad-scale patterns,drivers, and consequences of phenology

Plant phenology research has surged in recent decades, in part due to interest in phenological sensitivity to climate change and the vital role phenology plays in ecology. Many local-scale studies have generated important findings regarding the physiology, responses, and risks associated with shifts in plant phenology. By comparison, our understanding of regional- and global-scale phenology has been largely limited to remote sensing of green-up without the ability to differentiate among plant species. However, a new generation of analytical tools and data sources—including enhanced remote sensing products, digitized herbarium specimen data, and public participation in science—now permits investigating patterns and drivers of phenology across extensive taxonomic, temporal, and spatial scales, in an emerging field that we call macrophenology. Recent studies have highlighted how phenology affects dynamics at broad scales, including species interactions and ranges, carbon fluxes, and climate. At the cusp of this developing field of study, we review the theoretical and practical advances in four primary areas of plant macrophenology: (1) global patterns and shifts in plant phenology, (2) within-species changes in phenology as they mediate species' range limits and invasions at the regional scale, (3) broad-scale variation in phenology among species leading to ecological mismatches, and (4) interactions between phenology and global ecosystem processes. To stimulate future research, we describe opportunities for macrophenology to address grand challenges in each of these research areas, as well as recently available data sources that enhance and enable macrophenology research.     

Forecasting phenology under global warming

As a consequence of warming temperatures around the world, spring and autumn phenologies have been shifting, with corresponding changes in the length of the growing season. Our understanding of the spatial and interspecific variation of these changes, however, is limited. Not all species are responding similarly, and there is significant spatial variation in responses even within species. This spatial and interspecific variation complicates efforts to predict phenological responses to ongoing climate change, but must be incorporated in order to build reliable forecasts. Here, we use a long-term dataset (1953–2005) of plant phenological events in spring (flowering and leaf out) and autumn (leaf colouring and leaf fall) throughout Japan and South Korea to build forecasts that account for these sources of variability. Specifically, we used hierarchical models to incorporate the spatial variability in phenological responses to temperature to then forecast species'' overall and site-specific responses to global warming. We found that for most species, spring phenology is advancing and autumn phenology is getting later, with the timing of events changing more quickly in autumn compared with the spring. Temporal trends and phenological responses to temperature in East Asia contrasted with results from comparable studies in Europe, where spring events are changing more rapidly than are autumn events. Our results emphasize the need to study multiple species at many sites to understand and forecast regional changes in phenology.     

Adaptation, niche conservatism, and convergence: comparative studies of leaf evolution in the California chaparral

Small leaves and low specific leaf area (SLA) have long been viewed as adaptations to Mediterranean-type climates in many species of evergreen woody plants. However, paleobotanical and floristic evidence suggests that in many cases these traits originated prior to the advent of the summer-drought climate regime. In this study, molecular phylogenies and ancestral state reconstructions were used to test the hypothesis of adaptive leaf evolution in 12 lineages of evergreen shrubs in the California chaparral. Across all lineages there was a small but significant shift toward lower SLA, but there were no trends in leaf size evolution. For individual lineages, adaptive changes were detected in only three cases for SLA and in one case for leaf size. Three of these cases of evolutionary change were observed in taxa derived from cool temperate ancestors (e.g., Heteromeles). In contrast, most lineages originating from subtropical ancestors exhibited relative stasis in leaf trait evolution (e.g., Ceanothus). The absence of change suggests that ancestors of chaparral taxa had already acquired appropriate traits that contributed to their success under Mediterranean-type climates. These results illustrate how biogeographic history may influence patterns of trait evolution and adaptation and highlight the contribution of ecological sorting processes to the assembly and functional ecology of regional biotas.     

亚热带常绿阔叶林和暖温带落叶阔叶林叶片热值比较研究

植物干重热值(GCV)是衡量植物生命活动及组成成分的重要指标之一,反映了植物光合作用中固定太阳辐射的能力。利用氧弹量热仪测定了亚热带和暖温带两个典型森林生态系统常见的276种常见植物叶片的干重热值,探讨了亚热带和暖温带植物热值分布特征,以及不同生活型、乔木类型间植物热值的变化规律。实验结果发现:亚热带常绿阔叶林和暖温带落叶阔叶林叶片热值的平均值分别为17.83 k J/g(n=191)和17.21k J/g(n=85),整体表现为亚热带植物暖温带植物。不同地带性植被的植物叶片热值在不同生活型间表现出相似的规律,其中亚热带常绿阔叶林表现为:乔木(19.09 k J/g)灌木(17.87 k J/g)草本(16.65 k J/g);暖温带落叶阔叶林表现为:乔木(18.41 k J/g)灌木(17.94 k J/g)草本(16.53 k J/g);不同乔木类型间均呈现常绿乔木落叶乔木、针叶乔木阔叶乔木的趋势。落叶阔叶乔木表现为亚热带暖温带,而常绿针叶乔木则呈现亚热带暖温带的趋势。此外,我们对于两个分布区域内的4种针叶树种叶片热值进行了比较,发现华北落叶松(19.32 k J/g,暖温带)杉木(19.40 k J/g,亚热带)马尾松(19.82 k J/g,亚热带)油松(20.95 k J/g,暖温带)。亚热带常绿阔叶林和暖温带落叶阔叶林植物热值的特征及其变化规律,为森林生态系统的能量流动提供了理论基础。     

New insights on plant phenological response to temperature revealed from long‐term widespread observations in China

Constraints of temperature on spring plant phenology are closely related to plant growth, vegetation dynamics, and ecosystem carbon cycle. However, the effects of temperature on leaf onset, especially for winter chilling, are still not well understood. Using long‐term, widespread in situ phenology observations collected over China for multiple plant species, this study analyzes the quantitative response of leaf onset to temperature, and compares empirical findings with existing theories and modeling approaches, as implemented in 18 phenology algorithms. Results show that the growing degree days (GDD) required for leaf onset vary distinctly among plant species and geographical locations as well as at organizational levels (species and community), pointing to diverse adaptation strategies. Chilling durations (CHD) needed for releasing bud dormancy decline monotonously from cold to warm areas with very limited interspecies variations. Results also reveal that winter chilling is a crucial component of phenology models, and its effect is better captured with an index that accounts for the inhomogeneous effectiveness of low temperature to chilling rate than with the conventional CHD index. The impact of spring warming on leaf onset is nonlinear, better represented by a logistical function of temperature than by the linear function currently implemented in biosphere models. The optimized base temperatures for thermal accumulation and the optimal chilling temperatures are species‐dependent and average at 6.9 and 0.2°C, respectively. Overall, plants’ chilling requirement is not a constant, and more chilling generally results in less requirement of thermal accumulation for leaf onset. Our results clearly demonstrate multiple deficiencies of the parameters (e.g., base temperature) and algorithms (e.g., method for calculating GDD) in conventional phenology models to represent leaf onset. Therefore, this study not only advances our mechanistic and quantitative understanding of temperature controls on leaf onset but also provides critical information for improving existing phenology models.     

Spatial and temporal variations in the timing of leaf replacement in a Quercus cornelius‐mulleri population

Abstract. The phenology of spring leaf replacement was studied in a population of 46 evergreen scrub oaks (Quercus cornelius‐mulleri) at the edge of the Mojave Desert in each of five years over the period 1990–2001. The oaks occupied a site that spanned rocky slopes to sandy bajadas. The site receives variable annual rainfall (estimated 12‐yr average 195 mm; range in study years 67–706 mm). The spatial coordinates of all individuals were recorded, and in April, when leaf replacement was underway, individual replacement phenologies were assessed. Shrub sizes were recorded in three separate years, and in 2001 water potentials were measured. Individuals vary greatly in their timing of leaf replacement within years, and also between years. Many individuals with an early phenology one year are significantly later in the following year, and vice versa. While we detected weak influences on leaf replacement phenology due to shrub size, position within the site, and a genetic component, stronger influences were attributable to the phenology of the shrub in prior years, and to the phenology of neighbours within years. Neighbouring individuals that are close and/or large are significantly disparate in phenology, with one early and the other late. A potential mechanism of local resource depletion associated with costs to an early phenology is discussed.     

Multiscale modeling of spring phenology across Deciduous Forests in the Eastern United States

Phenological events, such as bud burst, are strongly linked to ecosystem processes in temperate deciduous forests. However, the exact nature and magnitude of how seasonal and interannual variation in air temperatures influence phenology is poorly understood, and model‐based phenology representations fail to capture local‐ to regional‐scale variability arising from differences in species composition. In this paper, we use a combination of surface meteorological data, species composition maps, remote sensing, and ground‐based observations to estimate models that better represent how community‐level species composition affects the phenological response of deciduous broadleaf forests to climate forcing at spatial scales that are typically used in ecosystem models. Using time series of canopy greenness from repeat digital photography, citizen science data from the USA National Phenology Network, and satellite remote sensing‐based observations of phenology, we estimated and tested models that predict the timing of spring leaf emergence across five different deciduous broadleaf forest types in the eastern United States. Specifically, we evaluated two different approaches: (i) using species‐specific models in combination with species composition information to ‘upscale’ model predictions and (ii) using repeat digital photography of forest canopies that observe and integrate the phenological behavior of multiple representative species at each camera site to calibrate a single model for all deciduous broadleaf forests. Our results demonstrate variability in cumulative forcing requirements and photoperiod cues across species and forest types, and show how community composition influences phenological dynamics over large areas. At the same time, the response of different species to spatial and interannual variation in weather is, under the current climate regime, sufficiently similar that the generic deciduous forest model based on repeat digital photography performed comparably to the upscaled species‐specific models. More generally, results from this analysis demonstrate how in situ observation networks and remote sensing data can be used to synergistically calibrate and assess regional parameterizations of phenology in models.     

Leveraging plant hydraulics to yield predictive and dynamic plant leaf allocation in vegetation models with climate change

Plant functional traits provide a link in process‐based vegetation models between plant‐level physiology and ecosystem‐level responses. Recent advances in physiological understanding and computational efficiency have allowed for the incorporation of plant hydraulic processes in large‐scale vegetation models. However, a more mechanistic representation of water limitation that determines ecosystem responses to plant water stress necessitates a re‐evaluation of trait‐based constraints for plant carbon allocation, particularly allocation to leaf area. In this review, we examine model representations of plant allocation to leaves, which is often empirically set by plant functional type‐specific allometric relationships. We analyze the evolution of the representation of leaf allocation in models of different scales and complexities. We show the impacts of leaf allocation strategy on plant carbon uptake in the context of recent advancements in modeling hydraulic processes. Finally, we posit that deriving allometry from first principles using mechanistic hydraulic processes is possible and should become standard practice, rather than using prescribed allometries. The representation of allocation as an emergent property of scarce resource constraints is likely to be critical to representing how global change processes impact future ecosystem dynamics and carbon fluxes and may reduce the number of poorly constrained parameters in vegetation models.     

It is about time: genetic variation in the timing of leaf‐litter inputs influences aquatic ecosystems

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Microbial effects on plant phenology and fitness

Plant development and the timing of developmental events (phenology) are tightly coupled with plant fitness. A variety of internal and external factors determine the timing and fitness consequences of these life-history transitions. Microbes interact with plants throughout their life history and impact host phenology. This review summarizes current mechanistic and theoretical knowledge surrounding microbe-driven changes in plant phenology. Overall, there are examples of microbes impacting every phenological transition. While most studies have focused on flowering time, microbial effects remain important for host survival and fitness across all phenological phases. Microbe-mediated changes in nutrient acquisition and phytohormone signaling can release plants from stressful conditions and alter plant stress responses inducing shifts in developmental events. The frequency and direction of phenological effects appear to be partly determined by the lifestyle and the underlying nature of a plant–microbe interaction (i.e., mutualistic or pathogenic), in addition to the taxonomic group of the microbe (fungi vs. bacteria). Finally, we highlight biases, gaps in knowledge, and future directions. This biotic source of plasticity for plant adaptation will serve an important role in sustaining plant biodiversity and managing agriculture under the pressures of climate change.     

热带森林植物功能群及其动态研究进展

热带森林极高的物种多样性和结构复杂性给生态学研究带来了很多挑战。植物功能群是对特定环境响应相似或对主要的生态过程具有相似作用的物种组合。应用植物功能群的方法是有效减少热带森林群落复杂性,并揭示其格局和过程的良好途径。在介绍植物功能群概念和划分途径的基础上,分析了热带森林植物功能群的时空动态规律。一般来讲,划分植物功能群通常有3种途径,并可通过5个步骤来完成。热带森林植物功能群的空间分布常受景观格局的制约,而环境异质性往往是影响不同植物功能群组配比例变化的直接原因。不同类型的植物功能群随演替过程发生显著的替代,而干扰体系和全球气候变化对功能群的动态过程具有重要的驱动作用。以功能群为基础的动态模型在模拟热带林群落动态和预测植被潜在分布等方面具有广阔的发展前景。探索有效的植物功能分类方法、构建完善的植物功能性状数据库、开发更为精确的功能群动态模型以及加强基于景观水平的植物功能群动态机制的认识等是未来热带森林植物功能群及其动态研究的重要方向。     

Testing a growth efficiency hypothesis with continental-scale phenological variations of common and cloned plants

Variation in the timing of plant phenology caused by phenotypic plasticity is a sensitive measure of how organisms respond to weather and climate variability. Although continental-scale gradients in climate and consequential patterns in plant phenology are well recognized, the contribution of underlying genotypic difference to the geography of phenology is less well understood. We hypothesize that different temperate plant genotypes require varying amount of heat energy for resuming annual growth and reproduction as a result of adaptation and other ecological and evolutionary processes along climatic gradients. In particular, at least for some species, the growing degree days (GDD) needed to trigger the same spring phenology events (e.g., budburst and flower bloom) may be less for individuals originated from colder climates than those from warmer climates. This variable intrinsic heat energy requirement in plants can be characterized by the term growth efficiency and is quantitatively reflected in the timing of phenophases—earlier timing indicates higher efficiency (i.e., less heat energy needed to trigger phenophase transitions) and vice versa compared to a standard reference (i.e., either a uniform climate or a uniform genotype). In this study, we tested our hypothesis by comparing variations of budburst and bloom timing of two widely documented plants from the USA National Phenology Network (i.e., red maple-Acer rubrum and forsythia-Forsythia spp.) with cloned indicator plants (lilac-Syringa x chinensis ‘Red Rothomagensis’) at multiple eastern US sites. Our results indicate that across the accumulated temperature gradient, the two non-clonal plants showed significantly more gradual changes than the cloned plants, manifested by earlier phenology in colder climates and later phenology in warmer climates relative to the baseline clone phenological response. This finding provides initial evidence supporting the growth efficiency hypothesis, and suggests more work is warranted. More studies investigating genotype-determined phenological variations will be useful for better understanding and prediction of the continental-scale patterns of biospheric responses to climate change.     

Spring leaf phenology and the diurnal temperature range in a temperate maple forest

Spring leaf phenology in temperate climates is intricately related to numerous aspects of the lower atmosphere [e.g., surface energy balance, carbon flux, humidity, the diurnal temperature range (DTR)]. To further develop and improve the accuracy of ecosystem and climate models, additional investigations of the specific nature of the relationships between spring leaf phenology and various ecosystem and climate processes are required in different environments. This study used visual observations of maple leaf phenology, below-canopy light intensities, and micrometeorological data collected during the spring seasons of 2008, 2009, and 2010 to examine the potential influence of leaf phenology on a seasonal transition in the trend of the DTR. The timing of a reversal in the DTR trend occurred near the time when the leaves were unfolding and expanding. The results suggest that the spring decline in the DTR can be attributed primarily to the effect of canopy closure on daily maximum temperature. These findings improve our understanding of the relationship between leaf phenology and the diurnal temperature range in temperate maple forests during the spring. They also demonstrate the necessity of incorporating accurate phenological data into ecosystem and climate models and warrant a careful examination of the extent to which canopy phenology is currently incorporated into existing models.     

The phenology of leaf quality and its within‐canopy variation is essential for accurate modeling of photosynthesis in tropical evergreen forests

Leaf quantity (i.e., canopy leaf area index, LAI), quality (i.e., per‐area photosynthetic capacity), and longevity all influence the photosynthetic seasonality of tropical evergreen forests. However, these components of tropical leaf phenology are poorly represented in most terrestrial biosphere models (TBMs). Here, we explored alternative options for the representation of leaf phenology effects in TBMs that employ the Farquahar, von Caemmerer & Berry (FvCB) representation of CO₂ assimilation. We developed a two‐fraction leaf (sun and shade), two‐layer canopy (upper and lower) photosynthesis model to evaluate different modeling approaches and assessed three components of phenological variations (i.e., leaf quantity, quality, and within‐canopy variation in leaf longevity). Our model was driven by the prescribed seasonality of leaf quantity and quality derived from ground‐based measurements within an Amazonian evergreen forest. Modeled photosynthetic seasonality was not sensitive to leaf quantity, but was highly sensitive to leaf quality and its vertical distribution within the canopy, with markedly more sensitivity to upper canopy leaf quality. This is because light absorption in tropical canopies is near maximal for the entire year, implying that seasonal changes in LAI have little impact on total canopy light absorption; and because leaf quality has a greater effect on photosynthesis of sunlit leaves than light limited, shade leaves and sunlit foliage are more abundant in the upper canopy. Our two‐fraction leaf, two‐layer canopy model, which accounted for all three phenological components, was able to simulate photosynthetic seasonality, explaining ~90% of the average seasonal variation in eddy covariance‐derived CO₂ assimilation. This work identifies a parsimonious approach for representing tropical evergreen forest photosynthetic seasonality in TBMs that utilize the FvCB model of CO₂ assimilation and highlights the importance of incorporating more realistic phenological mechanisms in models that seek to improve the projection of future carbon dynamics in tropical evergreen forests.     

Generalized additive mixed models for disentangling long‐term trends,local anomalies,and seasonality in fruit tree phenology

Quantifying temporal patterns of ephemeral plant structures such as leaves, flowers, and fruits gives insight into both plant and animal ecology. Different scales of temporal changes in fruits, for example within‐ versus across‐year variability, are driven by different processes, but are not always easy to disentangle. We apply generalized additive mixed models (GAMMs) to study a long‐term fruit presence–absence data set of individual trees collected from a high‐altitude Afromontane tropical rain forest site within Bwindi Impenetrable National Park (BINP), Uganda. Our primary aim was to highlight and evaluate GAMM methodology, and quantify both intra‐ and interannual changes in fruit production. First, we conduct several simulation experiments to study the practical utility of model selection and smooth term estimation relevant for disentangling intra‐ and interannual variability. These simulations indicate that estimation of nonlinearity and seasonality is generally accurately identified using asymptotic theory. Applied to the empirical data set, we found that the forest‐level fruiting variability arises from both regular seasonality and significant interannual variability, with the years 2009–2010 in particular showing a significant increase in the presence of fruits‐driven by increased productivity of most species, and a regular annual peak associated occurring at the end of one of the two dry seasons. Our analyses illustrate a statistical framework for disentangling short‐term increases/decreases in fruiting effort while pinpointing specific times in which fruiting is atypical, providing a first step for assessing the impacts of regular and irregular (e.g., climate change) abiotic covariates on fruiting phenology. Some consequences of the rich diversity of fruiting patterns observed here for the population biology of frugivores in BINP are also discussed.     

Landscape controls on the timing of spring,autumn, and growing season length in mid‐Atlantic forests

The timing of spring leaf development, trajectories of summer leaf area, and the timing of autumn senescence have profound impacts to the water, carbon, and energy balance of ecosystems, and are likely influenced by global climate change. Limited field‐based and remote‐sensing observations have suggested complex spatial patterns related to geographic features that influence climate. However, much of this variability occurs at spatial scales that inhibit a detailed understanding of even the dominant drivers. Recognizing these limitations, we used nonlinear inverse modeling of medium‐resolution remote sensing data, organized by day of year, to explore the influence of climate‐related landscape factors on the timing of spring and autumn leaf‐area trajectories in mid‐Atlantic, USA forests. We also examined the extent to which declining summer greenness (greendown) degrades the precision and accuracy of observations of autumn offset of greenness. Of the dominant drivers of landscape phenology, elevation was the strongest, explaining up to 70% of the spatial variation in the onset of greenness. Urban land cover was second in importance, influencing spring onset and autumn offset to a distance of 32 km from large cities. Distance to tidal water also influenced phenological timing, but only within ~5 km of shorelines. Additionally, we observed that (i) growing season length unexpectedly increases with increasing elevation at elevations below 275 m; (ii) along gradients in urban land cover, timing of autumn offset has a stronger effect on growing season length than does timing of spring onset; and (iii) summer greendown introduces bias and uncertainty into observations of the autumn offset of greenness. These results demonstrate the power of medium grain analyses of landscape‐scale phenology for understanding environmental controls on growing season length, and predicting how these might be affected by climate change.     

Predicting vegetation type through physiological and environmental interactions with leaf traits: evergreen and deciduous forests in an earth system modeling framework

Earth system models are incorporating plant trait diversity into their land components to better predict vegetation dynamics in a changing climate. However, extant plant trait distributions will not allow extrapolations to novel community assemblages in future climates, which will require a mechanistic understanding of the trade‐offs that determine trait diversity. In this study, we show how physiological trade‐offs involving leaf mass per unit area (LMA), leaf lifespan, leaf nitrogen, and leaf respiration may explain the distribution patterns of evergreen and deciduous trees in the temperate and boreal zones based on (1) an evolutionary analysis of a simple mathematical model and (2) simulation experiments of an individual‐based dynamic vegetation model (i.e., LM3‐PPA). The evolutionary analysis shows that these leaf traits set up a trade‐off between carbon‐ and nitrogen‐use efficiency at the scale of individual trees and therefore determine competitively dominant leaf strategies. As soil nitrogen availability increases, the dominant leaf strategy switches from one that is high in nitrogen‐use efficiency to one that is high in carbon‐use efficiency or, equivalently, from high‐LMA/long‐lived leaves (i.e., evergreen) to low‐LMA/short‐lived leaves (i.e., deciduous). In a region of intermediate soil nitrogen availability, the dominant leaf strategy may be either deciduous or evergreen depending on the initial conditions of plant trait abundance (i.e., founder controlled) due to feedbacks of leaf traits on soil nitrogen mineralization through litter quality. Simulated successional patterns by LM3‐PPA from the leaf physiological trade‐offs are consistent with observed successional dynamics of evergreen and deciduous forests at three sites spanning the temperate to boreal zones.