Amazon forest carbon dynamics predicted by profiles of canopy leaf area and light environment

Tropical forest structural variation across heterogeneous landscapes may control above‐ground carbon dynamics. We tested the hypothesis that canopy structure (leaf area and light availability) – remotely estimated from LiDAR – control variation in above‐ground coarse wood production (biomass growth). Using a statistical model, these factors predicted biomass growth across tree size classes in forest near Manaus, Brazil. The same statistical model, with no parameterisation change but driven by different observed canopy structure, predicted the higher productivity of a site 500 km east. Gap fraction and a metric of vegetation vertical extent and evenness also predicted biomass gains and losses for one‐hectare plots. Despite significant site differences in canopy structure and carbon dynamics, the relation between biomass growth and light fell on a unifying curve. This supported our hypothesis, suggesting that knowledge of canopy structure can explain variation in biomass growth over tropical landscapes and improve understanding of ecosystem function.     

Age‐related Crown Thinning in Tropical Forest Trees

Gap dynamics theory proposes that treefall gaps provide high light levels needed for regeneration in the understory, and by increasing heterogeneity in the light environment allow light‐demanding tree species to persist in the community. Recent studies have demonstrated age‐related declines in leaf area index of individual temperate trees, highlighting a mechanism for gradual changes in the forest canopy that may also be an important, but less obvious, driver of forest dynamics. We assessed the prevalence of age‐related crown thinning among 12 tropical canopy tree species sampled in lowland forests in Panama and Puerto Rico (total N = 881). Canopy gap fraction of individual canopy tree crowns was positively related to stem diameter at 1.3 m (diameter at breast height) in a pooled analysis, with 10 of 12 species showing a positive trend. Considered individually, a positive correlation between stem diameter and canopy gap fraction was statistically significant in 4 of 12 species, all of which were large‐statured canopy to emergent species: Beilschmiedia pendula, Ceiba pentandra, Jacaranda copaia, and Prioria copaifera. Pooled analyses also showed a negative relationship between liana abundance and canopy gap fraction, suggesting that lianas could be partially obscuring age‐related crown thinning. We conclude that age‐related crown thinning occurs in tropical forests, and could thus influence patterns of tree regeneration and tropical forest community dynamics.     

Short‐term Dynamics and the Effects of Biotic and Abiotic Factors on Plant Physiognomic Groups in a Hurricane‐impacted Lower Montane Tropical Forest

We assessed the short‐term effects of biotic (density, plant size) and abiotic factors (light), on the dynamics of physiognomically different plant groups (palms, tree ferns, lianas, and trees) in a hurricane‐impacted tropical wet montane forest, John Crow Mountains, Jamaica. All plants ≥2 cm (dbh) found within 45, 25 × 25 m permanent sample plots (2.8125 ha), established according to a randomized block design along an elevation gradient, were tagged and measured (dbh) in 2006 and re‐assessed in 2012 after Hurricane Dean (2007). Hemispheric light was measured in 2007 and 2008. Tree and liana size class distributions changed due to high mortality in the smallest size classes and their densities declined; however, palm and tree fern density remained unchanged. The dynamics of trees were only related to tree fern and liana dynamics (e.g., tree mortality was negatively related to liana recruitment etc.). Although pre‐ and posthurricane light was related to palm density and the density of the other plant groups, respectively, there were no significant changes in light. Tree survivorship increased with increasing dbh while posthurricane light and overall density influenced the growth and survivorship of tree species. Species importance value did not change, suggesting that direct regeneration may be the model of forest recovery following this small‐scale disturbance. Over the short term, tree species showed life history trade‐offs that aid species coexistence after this moderate/low disturbance event. Our study highlights that hurricanes with low impacts can have differential short‐ and possibly long‐term effects on different plant groups.     

山西灵空山天然松栎混交林群落特征与冠层结构

以2011年建设的山西灵空山4 hm²天然松栎混交林森林动态监测样地为研究平台,以400个10 m×10 m样方为测量单元,于2016年进行群落特征研究,采用半球面影像法(DHP)分析冠层结构和林下光照特征.结果表明: 样地内共有乔木5558株,共计25种,分属于10科15属.冠层开阔度(CO)集中在15.0%～25.0%,叶面积指数(LAI)集中在1.5～2.5,林下光环境参数集中在10.0%～30.0%.建群种在样地内的分布对冠层结构和林下光环境影响显著;冠层结构对林下光环境所有参数的影响方向一致,其中采用叶面积指数评价冠层结构动态的效果更佳;冠层开阔度和叶面积指数对林下光环境产生相反的影响,且均对散射光入射率影响程度最大.温性松栎混交林的林冠层整体较为均匀,林下光分布较为集中,林分树种组成与冠层结构对林下光照影响显著.     

中亚热带森林单木地上生物量的机载激光雷达估测

以雪峰山武冈林场为研究对象,利用遥感数据和地面实测样地数据,研究机载激光雷达（LiDAR）估测中亚热带森林乔木层单木地上生物量的能力.利用条件随机场和最优化方法实现LiDAR点云的单木分割,以单木尺度为对象提取的植被点云空间结构、回波特征以及地形特征等作为遥感变量,采用回归模型估测乔木层地上生物量.结果表明: 针叶林、阔叶林和针阔混交林的单木识别率分别为93%、86%和60%;多元逐步回归模型的调整决定系数分别为0.83、0.81和0.74,均方根误差分别为28.22、29.79和32.31 t·hm^-2;以冠层体积、树高百分位值、坡度和回波强度值构成的模型精度明显高于以树高为因子的传统回归模型精度.以单木为对象从LiDAR点云中提取的遥感变量有助于提高森林生物量估测精度.
     

Prediction of forest aboveground net primary production from high‐resolution vertical leaf‐area profiles

Temperature and precipitation explain about half the variation in aboveground net primary production (ANPP) among tropical forest sites, but determinants of remaining variation are poorly understood. Here, we test the hypothesis that the amount of leaf area, and its vertical arrangement, predicts ANPP when other variables are held constant. Using measurements from airborne lidar in a lowland Neotropical rain forest, we quantify vertical leaf‐area profiles and develop models of ANPP driven by leaf area and other measurements of forest structure. Vertical leaf‐area profiles predict 38% of the variation among plots. This number is 4.5 times greater than models using total leaf area (disregarding vertical arrangement) and 2.1 times greater than models using canopy height alone. Furthermore, ANPP predictions from vertical leaf‐area profiles were less biased than alternate metrics. Variation in ANPP not attributable to temperature or precipitation can be predicted by the vertical distribution of leaf area in this system.     

Evidence of non‐stationary relationships between climate and forest responses: Increased sensitivity to climate change in Iberian forests

Climate and forest structure are considered major drivers of forest demography and productivity. However, recent evidence suggests that the relationships between climate and tree growth are generally non‐stationary (i.e. non‐time stable), and it remains uncertain whether the relationships between climate, forest structure, demography and productivity are stationary or are being altered by recent climatic and structural changes. Here we analysed three surveys from the Spanish Forest Inventory covering c. 30 years of information and we applied mixed and structural equation models to assess temporal trends in forest structure (stand density, basal area, tree size and tree size inequality), forest demography (ingrowth, growth and mortality) and above‐ground forest productivity. We also quantified whether the interactive effects of climate and forest structure on forest demography and above‐ground forest productivity were stationary over two consecutive time periods. Since the 1980s, density, basal area and tree size increased in Iberian forests, and tree size inequality decreased. In addition, we observed reductions in ingrowth and growth, and increases in mortality. Initial forest structure and water availability mainly modulated the temporal trends in forest structure and demography. The magnitude and direction of the interactive effects of climate and forest structure on forest demography changed over the two time periods analysed indicating non‐stationary relationships between climate, forest structure and demography. Above‐ground forest productivity increased due to a positive balance between ingrowth, growth and mortality. Despite increasing productivity over time, we observed an aggravation of the negative effects of climate change and increased competition on forest demography, reducing ingrowth and growth, and increasing mortality. Interestingly, our results suggest that the negative effects of climate change on forest demography could be ameliorated through forest management, which has profound implications for forest adaptation to climate change.     

Contrasting patterns of tree mortality in late‐successional Picea abies stands in two areas in northern Fennoscandia

Question: What are tree mortality rates and how and why do they vary in late‐successional Picea abies‐dominated forests? Do observed tree mortality patterns allow comparative assessment of models of long‐term stand development? Location: Northern boreal Fennoscandia. Methods: We measured stand structure in 10 stands in two different areas. We determined age distributions and constructed a chronology of tree deaths by cross‐dating the years of death of randomly sampled dead trees. Results: The stands in the two areas had contrasting tree age distributions, despite similar live tree structure. In one area, stands were relatively even‐aged and originated following a stand‐replacing fire 317 years earlier. The stands in the second area had an uneven age structure and virtually no signs of past fires, suggesting a very long period since the last major disturbance. The younger stands were characterized by a high mortality rate and inter‐annual variation, which we attributed to senescence of the relatively even‐aged stands approaching the maximum age of P. abies. In contrast, the tree mortality rates in the older stands were low and relatively stable. Conclusions: Patterns of tree mortality were, to a large extent, dependent on the time since the last stand‐replacing disturbance, suggesting that northern boreal P. abies stands eventually reach a shifting mosaic state maintained through small‐scale dynamics, but the time needed to reach this state appears to be lengthy; even 300 years after a forest fire stands showed changes in patterns of tree mortality that were related to the developmental stage of the stands.     

粤北山地常绿阔叶林自然干扰后冠层结构与林下光照动态

以粤北车八岭2008年受冰灾破坏的山地常绿阔叶林为研究对象,设置2 hm2固定样地开展连续3a(2008—2010年)的群落调查,并采用半球面影像技术(Hemispherical photography)获取冠层结构和林下光照指标,分析灾后森林演替过程中冠层结构和林下光照的动态。研究发现:1)灾后森林恢复过程中,样地林下光照(直射光、散射光和总光照)均随林冠开度的减少、叶面积指数的增加而减少;2)从2008到2010年,各年度冠层结构和林下光照的差异均极显著(P<0.0001),但年间差异程度有逐年减少的趋势;3)灾后森林恢复前3a,林下直射光对林下总光照的贡献大于散射光,其时空波动性也大于散射光;4)林冠开度对冠层结构的反映程度比叶面积指数高,冠层结构对林下散射光的影响比对直射光大。灾后林木先是迅速生长然后生长速度缓慢下来并逐渐稳定,随森林逐渐郁闭林下光照也随之减少,其中林冠开度用于评价冠层结构动态的效果更佳,林下直射光比散射光的时空变化更复杂。     

Modelling canopy CO2 fluxes: are ‘big‐leaf’ simplifications justified?

• 1 The ‘big‐leaf’ approach to calculating the carbon balance of plant canopies is evaluated for inclusion in the ETEMA model framework. This approach assumes that canopy carbon fluxes have the same relative responses to the environment as any single leaf, and that the scaling from leaf to canopy is therefore linear.
• 2 A series of model simulations was performed with two models of leaf photosynthesis, three distributions of canopy nitrogen, and two levels of canopy radiation detail. Leaf‐ and canopy‐level responses to light and nitrogen, both as instantaneous rates and daily integrals, are presented.
• 3 Observed leaf nitrogen contents of unshaded leaves are over 40% lower than the big‐leaf approach requires. Scaling from these leaves to the canopy using the big‐leaf approach may underestimate canopy photosynthesis by ~20%. A leaf photosynthesis model that treats within‐leaf light extinction displays characteristics that contradict the big‐leaf theory. Observed distributions of canopy nitrogen are closer to those required to optimize this model than the homogeneous model used in the big‐leaf approach.
• 4 It is theoretically consistent to use the big‐leaf approach with the homogeneous photosynthesis model to estimate canopy carbon fluxes if canopy nitrogen and leaf area are known and if the distribution of nitrogen is assumed optimal. However, real nitrogen profiles are not optimal for this photosynthesis model, and caution is necessary in using the big‐leaf approach to scale satellite estimates of leaf physiology to canopies. Accurate prediction of canopy carbon fluxes requires canopy nitrogen, leaf area, declining nitrogen with canopy depth, the heterogeneous model of leaf photosynthesis and the separation of sunlit and shaded leaves. The exact nitrogen profile is not critical, but realistic distributions can be predicted using a simple model of canopy nitrogen allocation.

     

Allometric constraints and competition enable the simulation of size structure and carbon fluxes in a dynamic vegetation model of tropical forests (LM3PPA‐TV)

Tropical forests are a key determinant of the functioning of the Earth system, but remain a major source of uncertainty in carbon cycle models and climate change projections. In this study, we present an updated land model (LM3PPA‐TV) to improve the representation of tropical forest structure and dynamics in Earth system models (ESMs). The development and parameterization of LM3PPA‐TV drew on extensive datasets on tropical tree traits and long‐term field censuses from Barro Colorado Island (BCI), Panama. The model defines a new plant functional type (PFT) based on the characteristics of shade‐tolerant, tropical tree species, implements a new growth allocation scheme based on realistic tree allometries, incorporates hydraulic constraints on biomass accumulation, and features a new compartment for tree branches and branch fall dynamics. Simulation experiments reproduced observed diurnal and seasonal patterns in stand‐level carbon and water fluxes, as well as mean canopy and understory tree growth rates, tree size distributions, and stand‐level biomass on BCI. Simulations at multiple sites captured considerable variation in biomass and size structure across the tropical forest biome, including observed responses to precipitation and temperature. Model experiments suggested a major role of water limitation in controlling geographic variation forest biomass and structure. However, the failure to simulate tropical forests under extreme conditions and the systematic underestimation of forest biomass in Paleotropical locations highlighted the need to incorporate variation in hydraulic traits and multiple PFTs that capture the distinct floristic composition across tropical domains. The continued pressure on tropical forests from global change demands models which are able to simulate alternative successional pathways and their pace to recovery. LM3PPA‐TV provides a tool to investigate geographic variation in tropical forests and a benchmark to continue improving the representation of tropical forests dynamics and their carbon storage potential in ESMs.     

Structural dynamics of a natural mixed deciduous forest in western Thailand

Abstract. Structural dynamics of a natural tropical seasonal – mixed deciduous – forest were studied over a 4-yr period at Mae Klong Watershed Research Station, Kanchanaburi Province, western Thailand, with particular reference to the role of forest fires and undergrowth bamboos. All trees > 5 cm DBH in a permanent plot of 200 m × 200 m were censused every two years from 1992 to 1996. The forest was characterized by a low stem density and basal area and relatively high species diversity. Both the bamboo undergrowth and frequent forest fires could be dominant factors that prevent continuous regeneration. Recruitment, mortality, gain (growth of survival tree plus ingrowth) and loss in basal area (by tree death) during the four years were 6.70%/yr, 2.91%/yr, 1.22%/yr and 1.34%/yr, respectively. Mortality was size dependent; middle size trees (30–50 cm) had the lowest mortality, while the smallest (5–10 cm) had the highest mortality. Tree recruitment was observed particularly in the first two years, mostly in the area where die-back of undergrowth bamboo occurred. The bias of the spatial distribution of recruitment to the area of bamboo die-back was significant and stronger than that to the forest canopy gaps. Successful regeneration of trees which survive competition with other herbs and trees after dieback of bamboo could occur when repeated forest fires did not occur in subsequent years. It is suggested that both the fire disturbance regime and bamboo life-cycle greatly influence the structure and dynamics of this seasonal tropical forest.     

Developing a Multilayered Integrated Numerical Model of Surface Physics – Growing Plants Interaction (MINoSGI)

This paper describes a Multilayered Integrated Numerical Model of Surface Physics – Growing Plants Interaction (MINoSGI), which represents interactions between the dynamics of forest ecosystems and microclimate. Aiming at a large‐scale study in the future, we describe forest dynamics by using area‐averaged prognostic equations for thedistributions of plant density and plant weight with respect to plant height classes, instead of individual‐based treatments for small‐scale forest patches. Growth and mortality of plants are modelled based on the carbon balance of each plant height class. The area‐averaged microclimate (e.g., light, wind speed, temperature, humidity, CO₂ concentration) within the forest canopy is simulated by a one‐dimensional multilayer canopy model, which includes most of the physical and physiological processes that control the forest microclimate. Owing to its multilayered framework, a direct specification is possible for the difference in the growing environment among plants of different size and species. Given hourly meteorological conditions, the model outputs energy, water, CO₂ and momentum fluxes to and from a forest, of which the structure changes through competition among plants. The model's performance was tested by comparing its outputs with observed data on the development of plant size distribution taken over a 5‐year period in an evergreen coniferous (Cryptomeria japonica) forest. The model produced realistic estimates of the total biomass increments during the period. The ratio of net primary production to gross primary production (=0.45) was consistent with previous estimates for temperate forests. The bimodal seasonal pattern in net ecosystem production was similar to the seasonal trend in the CO₂ flux measured over a forest of the same species. Although some limitations due to the one‐dimensional representation of microclimate were noticeable, the model adequately simulated distributions of annual growth rate, plant weight and diameter across plant height classes. Since the basic equations can be extended to include the effect of spatial variability with marginal increase of computational costs, the present model framework is feasible for large‐scale studies.     

Spatial and temporal dynamics of forest canopy gaps following selective logging in the eastern Amazon

Selective logging is a dominant form of land use in the Amazon basin and throughout the humid tropics, yet little is known about the spatial variability of forest canopy gap formation and closure following timber harvests. We established chronosequences of large‐area (14–158 ha) selective logging sites spanning a 3.5‐year period of forest regeneration and two distinct harvest methods: conventional logging (CL) and reduced‐impact logging (RIL). Our goals were to: (1) determine the spatial characteristics of canopy gap fraction immediately following selective logging in the eastern Amazon; (2) determine the degree and rate of canopy closure in early years following harvest among the major landscape features associated with logging – tree falls, roads, skid trails and log decks; and (3) quantify spatial and temporal differences in canopy opening and closure in high‐ and low‐damage harvests (CL vs. RIL). Across a wide range of harvest intensities (2.6–6.4 felled trees ha^?1), the majority of ground damage occurred as skid trails (4–12%), whereas log decks and roads were only a small contributor to the total ground damage (<2%). Despite similar timber harvest intensities, CL resulted in more ground damage than RIL. Neither the number of log decks nor their individual or total area was correlated with the number of trees removed or intensity of tree harvesting (trees ha^?1). The area of skids was well correlated with the ground area damaged (m²) per tree felled. In recently logged forest (0.5 years postharvest), gap fractions were highest in log decks (mean RIL=0.83, CL=0.99) and lowest in tree‐fall areas (RIL: 0.26, CL: 0.41). However, the small surface area of log decks made their contribution to the total area‐integrated forest gap fraction minor. In contrast, tree falls accounted for more than two‐thirds of the area disturbed, but the canopy gaps associated with felled trees were much smaller than for log decks, roads and skids. Canopy openings decreased in size with distance from each felled tree crown. At 0.5 years postharvest, the area initially affected by the felling of each tree was approximately 100 m in radius for CL and 50 m for RIL. Initial decreases in gap fraction during the first 1.5 years of regrowth diminished in subsequent years. Throughout the 3.5‐year period of forest recovery, tree‐fall gap fractions remained higher in CL than in RIL treatments, but canopy gap closure rates were higher in CL than in RIL areas. During the observed recovery period, the canopy gap area affected by harvesting decreased in radius around each felled tree from 100 to 40 m in CL, and from 50 to 10 m in RIL. The results suggest that the full spatial and temporal dynamics of canopy gap fraction must be understood and monitored to predict the effects of selective logging on regional energy balance and climate regimes, biogeochemical processes including carbon cycling, and plant and faunal population dynamics. This paper also shows that remote sensing of log decks alone will not provide an accurate assessment of total forest area impacted by selective logging, nor will it be closely correlated to damage levels and canopy gap closure rates.     

Climate variability drives recent tree mortality in Europe

Tree mortality is an important process in forest ecosystems, frequently hypothesized to be highly climate sensitive. Yet, tree death remains one of the least understood processes of forest dynamics. Recently, changes in tree mortality have been observed in forests around the globe, which could profoundly affect ecosystem functioning and services provisioning to society. We describe continental‐scale patterns of recent tree mortality from the only consistent pan‐European forest monitoring network, identifying recent mortality hotspots in southern and northern Europe. Analyzing 925,462 annual observations of 235,895 trees between 2000 and 2012, we determine the influence of climate variability and tree age on interannual variation in tree mortality using Cox proportional hazard models. Warm summers as well as high seasonal variability in precipitation increased the likelihood of tree death. However, our data also suggest that reduced cold‐induced mortality could compensate increased mortality related to peak temperatures in a warming climate. Besides climate variability, age was an important driver of tree mortality, with individual mortality probability decreasing with age over the first century of a trees life. A considerable portion of the observed variation in tree mortality could be explained by satellite‐derived net primary productivity, suggesting that widely available remote sensing products can be used as an early warning indicator of widespread tree mortality. Our findings advance the understanding of patterns of large‐scale tree mortality by demonstrating the influence of seasonal and diurnal climate variation, and highlight the potential of state‐of‐the‐art remote sensing to anticipate an increased likelihood of tree mortality in space and time.     

Leaf traits determine the growth-survival trade-off across rain forest tree species

A dominant hypothesis explaining tree species coexistence in tropical forest is that trade-offs in characters allow species to adapt to different light environments, but tests for this hypothesis are scarce. This study is the first that uses a theoretical plant growth model to link leaf trade-offs to whole-plant performances and to differential performances across species in different light environments. Using data of 50 sympatric tree species from a Bolivian rain forest, we observed that specific leaf area and photosynthetic capacity codetermined interspecific height growth variation in a forest gap; that leaf survival rate determined the variation in plant survival rate under a closed canopy; that predicted height growth and plant survival rate matched field observations; and that fast-growing species had low survival rates for both field and predicted values. These results show how leaf trade-offs influence differential tree performance and tree species' coexistence in a heterogeneous light environment.     

Structural overshoot of tree growth with climate variability and the global spectrum of drought‐induced forest dieback

Ongoing climate change poses significant threats to plant function and distribution. Increased temperatures and altered precipitation regimes amplify drought frequency and intensity, elevating plant stress and mortality. Large‐scale forest mortality events will have far‐reaching impacts on carbon and hydrological cycling, biodiversity, and ecosystem services. However, biogeographical theory and global vegetation models poorly represent recent forest die‐off patterns. Furthermore, as trees are sessile and long‐lived, their responses to climate extremes are substantially dependent on historical factors. We show that periods of favourable climatic and management conditions that facilitate abundant tree growth can lead to structural overshoot of aboveground tree biomass due to a subsequent temporal mismatch between water demand and availability. When environmental favourability declines, increases in water and temperature stress that are protracted, rapid, or both, drive a gradient of tree structural responses that can modify forest self‐thinning relationships. Responses ranging from premature leaf senescence and partial canopy dieback to whole‐tree mortality reduce canopy leaf area during the stress period and for a lagged recovery window thereafter. Such temporal mismatches of water requirements from availability can occur at local to regional scales throughout a species geographical range. As climate change projections predict large future fluctuations in both wet and dry conditions, we expect forests to become increasingly structurally mismatched to water availability and thus overbuilt during more stressful episodes. By accounting for the historical context of biomass development, our approach can explain previously problematic aspects of large‐scale forest mortality, such as why it can occur throughout the range of a species and yet still be locally highly variable, and why some events seem readily attributable to an ongoing drought while others do not. This refined understanding can facilitate better projections of structural overshoot responses, enabling improved prediction of changes in forest distribution and function from regional to global scales.     

Species-specific Seedling Responses to Hurricane Disturbance in a Puerto Rican Rain Forest1

Seedling dynamics were followed in a Puerto Rican forest for 20 months following a severe hurricane to study the interactive effects of hurricane debris, nutrients, and light on seedling diversity, density, growth, and mortality. Three treatments (debris removal, an unaltered control with hurricane debris, and chemical fertilization added to hurricane debris) altered levels of forest debris and soil nutrients. Canopy openness was measured twice using hemispherical photographs of the canopy. We examined the demographic responses of six common species to treatments over time. Seedling densities increased for all six species but the only significant treatment effects were increased densities of the pioneer tree Cecropia and the shrub Palicourea in the debris removal treatment. Seedling growth declined with declining light levels for four species but not for the pioneer tree Alchornea or the non‐pioneer tree Dacryodes. Only Cecropia and the non‐pioneer tree Chionanthus had treatment effects on growth. Mortality also differed among species and tended to be highest in the fertilized plots for all but Cecropia and Dacryodes. We found only some of the expected differences between pioneer and non‐pioneer plants, as each species had a unique response to the patchy distributions of organic debris, nutrients, and light following the hurricane. High local species diversity was maintained through the individualistic responses of seedlings after a disturbance.     

Impacts of a spring heat wave on canopy processes in a northern hardwood forest

Heat wave frequency, duration, and intensity are predicted to increase with global warming, but the potential impacts of short‐term high temperature events on forest functioning remain virtually unstudied. We examined canopy processes in a forest in Central Ontario following 3 days of record‐setting high temperatures (31–33 °C) that coincided with the peak in leaf expansion of dominant trees in late May 2010. Leaf area dynamics, leaf morphology, and leaf‐level gas‐exchange were compared to data from prior years of sampling (2002–2008) at the same site, focusing on Acer saccharum Marsh., the dominant tree in the region. Extensive shedding of partially expanded leaves was observed immediately following high temperature days, with A. saccharum losing ca. 25% of total leaf production but subsequently producing an unusual second flush of neoformed leaves. Both leaf losses and subsequent reflushing were highest in the upper canopy; however, retained preformed leaves and neoformed leaves showed reduced size, resulting in an overall decline in end‐of‐season leaf area index of 64% in A. saccharum, and 16% in the entire forest. Saplings showed lower leaf losses, but also a lower capacity to reflush relative to mature trees. Both surviving preformed and neoformed leaves had severely depressed photosynthetic capacity early in the summer of 2010, but largely regained photosynthetic competence by the end of the growing season. These results indicate that even short‐term heat waves can have severe impacts in northern forests, and suggest a particular vulnerability to high temperatures during the spring period of leaf expansion in temperate deciduous forests.     

An analytical model of stand dynamics as a function of tree growth, mortality and recruitment: the shade tolerance-stand structure hypothesis revisited

Light competition and interspecific differences in shade tolerance are considered key determinants of forest stand structure and dynamics. Specifically two main stand diameter distribution types as a function of shade tolerance have been proposed based on empirical observations. All-aged stands of shade tolerant species tend to have steeply descending, monotonic diameter distributions (inverse J-shaped curves). Shade intolerant species in contrast typically exhibit normal (unimodal) tree diameter distributions due to high mortality rates of smaller suppressed trees. In this study we explore the generality of this hypothesis which implies a causal relationship between light competition or shade tolerance and stand structure. For this purpose we formulate a partial differential equation system of stand dynamics as a function of individual tree growth, recruitment and mortality which allows us to explore possible individual-based mechanisms--e.g. light competition-underlying observed patterns of stand structure--e.g. unimodal or inverse J-shaped equilibrium diameter curves. We find that contrary to expectations interspecific differences in growth patterns can result alone in any of the two diameter distributions types observed in the field. In particular, slow growing species can present unimodal equilibrium curves even in the absence of light competition. Moreover, light competition and shade intolerance evaluated both at the tree growth and mortality stages did not have a significant impact on stand structure that tended to converge systematically towards an inverse J-shaped curves for most tree growth scenarios. Realistic transient stand dynamics for even aged stands of shade intolerant species (unimodal curves) were only obtained when recruitment was completely suppressed, providing further evidence on the critical role played by juvenile stages of tree development (e.g. the sampling stage) on final forest structure and composition. The results also point out the relevance of partial differential equations systems as a tool for exploring the individual-level mechanisms underpinning forest structure, particularly in relation to more complex forest simulation models that are more difficult to analyze and to interpret from a biological point of view.