Patterns and Drivers of Tree Mortality in Iberian Forests: Climatic Effects Are Modified by Competition

Tree mortality is a key process underlying forest dynamics and community assembly. Understanding how tree mortality is driven by simultaneous drivers is needed to evaluate potential effects of climate change on forest composition. Using repeat-measure information from c. 400,000 trees from the Spanish Forest Inventory, we quantified the relative importance of tree size, competition, climate and edaphic conditions on tree mortality of 11 species, and explored the combined effect of climate and competition. Tree mortality was affected by all of these multiple drivers, especially tree size and asymmetric competition, and strong interactions between climate and competition were found. All species showed L-shaped mortality patterns (i.e. showed decreasing mortality with tree size), but pines were more sensitive to asymmetric competition than broadleaved species. Among climatic variables, the negative effect of temperature on tree mortality was much larger than the effect of precipitation. Moreover, the effect of climate (mean annual temperature and annual precipitation) on tree mortality was aggravated at high competition levels for all species, but especially for broadleaved species. The significant interaction between climate and competition on tree mortality indicated that global change in Mediterranean regions, causing hotter and drier conditions and denser stands, could lead to profound effects on forest structure and composition. Therefore, to evaluate the potential effects of climatic change on tree mortality, forest structure must be considered, since two systems of similar composition but different structure could radically differ in their response to climatic conditions.     

The dynamics of hollowing in annually burnt savanna trees and its effect on adult tree mortality

Savanna trees often display significant hollows due to the combined action of fire and termites (N’Dri et al., J Trop Ecol 27:269–278, 2011). Observations have shown that internal cavities caused by termites in tree stems often result in external hollows under annual fire regimes, and it is quite possible that such hollows/fire interaction may increase the probability of trunk or big branch breakage and/or tree mortality. A study of individual tree and branch mortality as a function of prior termite and fire damage was undertaken in a West African savanna (Lamto, Côte d’Ivoire) where most of the trees naturally have hollows in their stems due to termite and fire interaction. Our goal was to examine the dynamics of hollowing and to determine whether hollowing significantly affected tree mortality. Branch and whole plant mortality were quantified for dominant tree species according to their initial hollow state and height. Four different responses were obtained depending on tree species: (1) mortality increased with cavity severity and tree size (Piliostigma thonningii), (2) mortality depended on tree size only (Bridelia ferruginea), (3) no mortality even after being hollowed by termites and externally damaged by fire (Crossopteryx febrifuga, the species with the highest proportion of individuals with hollows yet the greatest background survival time, 14 ± 2 years) and (4) high mortality, but few hollow trees suggesting a weak resistance to hollowing (Cussonia arborea which was insensitive to all the factors examined in this study). For species resistant to hollowing, tree mortality was rare; alternatively, for species prone to hollowing, whole trees died quickly and before the most severe hollow classes could be observed. Long-term demographic data yielded population-level mortality estimates of adult trees at least four times lower in fire-exclusion zones than that in fire-prone areas. Because hollow dynamics interact with fire in affecting adult mortality of some dominant tree species, fire management is important for a sustainable woody component of these savannas.     

Influences of forest structure, climate and species composition on tree mortality across the eastern US

Few studies have quantified regional variation in tree mortality, or explored whether species compositional changes or within-species variation are responsible for regional patterns, despite the fact that mortality has direct effects on the dynamics of woody biomass, species composition, stand structure, wood production and forest response to climate change. Using bayesian analysis of over 430,000 tree records from a large eastern US forest database we characterised tree mortality as a function of climate, soils, species and size (stem diameter). We found (1) mortality is U-shaped vs. stem diameter for all 21 species examined; (2) mortality is hump-shaped vs. plot basal area for most species; (3) geographical variation in mortality is substantial, and correlated with several environmental factors; and (4) individual species vary substantially from the combined average in the nature and magnitude of their mortality responses to environmental variation. Regional variation in mortality is therefore the product of variation in species composition combined with highly varied mortality-environment correlations within species. The results imply that variation in mortality is a crucial part of variation in the forest carbon cycle, such that including this variation in models of the global carbon cycle could significantly narrow uncertainty in climate change predictions.     

Rodent seed predation: effects on seed survival, recruitment, abundance, and dispersion of bird-dispersed tropical trees

Tropical tree species vary widely in their pattern of spatial dispersion. We focus on how seed predation may modify seed deposition patterns and affect the abundance and dispersion of adult trees in a tropical forest in India. Using plots across a range of seed densities, we examined whether seed predation levels by terrestrial rodents varied across six large-seeded, bird-dispersed tree species. Since inter-specific variation in density-dependent seed mortality may have downstream effects on recruitment and adult tree stages, we determined recruitment patterns close to and away from parent trees, along with adult tree abundance and dispersion patterns. Four species (Canarium resiniferum, Dysoxylum binectariferum, Horsfieldia kingii, and Prunus ceylanica) showed high predation levels (78.5-98.7%) and increased mortality with increasing seed density, while two species, Chisocheton cumingianus and Polyalthia simiarum, showed significantly lower seed predation levels and weak density-dependent mortality. The latter two species also had the highest recruitment near parent trees, with most abundant and aggregated adults. The four species that had high seed mortality had low recruitment under parent trees, were rare, and had more spaced adult tree dispersion. Biotic dispersal may be vital for species that suffer density-dependent mortality factors under parent trees. In tropical forests where large vertebrate seed dispersers but not seed predators are hunted, differences in seed vulnerability to rodent seed predation and density-dependent mortality can affect forest structure and composition.     

Predicting tree death for Fagus sylvatica and Abies alba using permanent plot data

Question: How well can mortality probabilities of deciduous trees(Fagus sylvatica) and conifers (Abies alba) be predicted using permanent plot data that describe growth patterns, tree species, tree size and site conditions? Location: Fagus forests in the montane belt of the Jura folds (Switzerland). Method: Permanent plot data were used to develop and validate logistic regression models predicting survival probabilities of individual trees. Backward model selection led to a reduced model containing the growth‐related variable ‘relative basal area increment’ (growth‐dependent mortality) and variables not directly reflecting growth such as species, size and site (growth‐independent mortality). Results: The growth‐mortality relationship was the same for both species (growth‐dependent mortality). However, species, site and tree size also influenced mortality probabilities (growth‐independent mortality). The predicted survival probabilities of the final model were well calibrated, and the model showed an excellent discriminatory power (area under the receiver operating characteristic curve = 0.896). Conclusion: Mortality probabilities of Fagus sylvatica and Abies alba can be predicted with high discriminatory power using a well calibrated logistic regression model. Extending this case study to a larger number of tree species and sites could provide species‐ and site‐specific tree mortality models that allow for more realistic projections of forest succession.     

Tree mortality of five major species on Hokkaido Island, northern Japan

The mortality rates of five major tree species (Abies sachalinensis, Acer mono, Magnolia obovata, Quercus crispula, Tilia japonica) on Hokkaido Island, northern Japan were modeled using logistic regressions based on the long-term observation of 8929 individuals in 65 permanent plots. Individual size, recent growth, one-sided and two-sided interactions with neighbors, climatic and topographic factors were used as independent variables in the logistic regressions, and relevant variables were selected using the stepwise method. For all species analyzed, the final models significantly explained tree mortality rates. The regression analyses indicated that individual size and/or recent growth had effects on tree mortality. A significant effect of one-sided interaction on tree mortality was detected for three species, and a significant effect of two-sided interaction was detected for two species. The interactions with neighbors were not necessarily competitive. For all species analyzed, climatic and topographic factors affected tree mortality. The mortality models obtained can be used for forest dynamics simulations. One ecological question was examined using these models. Species that can grow fast in forest stands with smaller basal areas tended to have high mortality rates in forest stands with larger basal areas. Some ecological characteristics of the analyzed species are described based on the results of the regressions.     

Climate change‐associated tree mortality increases without decreasing water availability

Temporal increases of tree mortality have been observed in regions where global warming has decreased long‐term water availability and/or induced droughts. However, temporal decreases in water availability are not a global phenomenon. Understanding how water deficit‐free forests respond to the recent effects of climate change is paramount towards a full appreciation of the impacts of climate change on global forests. Here, we reveal temporally increasing tree mortality across all study species over the last three decades in the central boreal forests of Canada, where long‐term water availability has increased without apparent climate change‐associated drought. In addition, we find that the effects of conspecific tree‐to‐tree competition have intensified temporally as a mechanism for the increased mortality of shade‐intolerant tree species. Our results suggest that the consequences of climate change on tree mortality are more profound than previously thought.     

Species‐ and site‐specific impacts of an invasive herbivore on tree survival in mixed forests

Invasive herbivores are often managed to limit their negative impact on plant populations, but herbivore density – plant damage relationships are notoriously spatially and temporally variable. Site and species characteristics (both plant and herbivore) must be considered when assessing the potential for herbivore damage, making it difficult to set thresholds for efficient management. Using the invasive brushtail possum Trichosurus vulpecula in New Zealand as a case study, we parameterized a generic model to predict annual probability of browse‐induced mortality of five tree species at 12 sites. We compared predicted and observed tree mortality for each species + site combination to establish herbivore abundance – tree mortality thresholds for each site on a single and combined tree species basis. Model results indicated it is likely that possum browse was the primary cause of all tree mortality at nine of the 12 species‐site combinations, allowing us to estimate site‐specific thresholds below which possum population numbers should be reduced and maintained to keep tree mortality under a predetermined level, for example 0.5% per year. The browse model can be used to set site‐ and species‐specific management action thresholds, and can be adapted easily for other plant or herbivore species. Results for multiple plant or herbivore species at a single site can be combined to create conservative, site‐wide management strategies, and used to: determine which sites will be affected most by changes in herbivore abundance; quantify thresholds for herbivore management; and justify expenditure on herbivore control.     

Effects of tree mortality on the spatial patterns and interspecific associations of plant species in a Quercus aliena var. acuteserrata forest in Baotianman,Henan, China

Aims Tree mortality is an important ecological process in forest ecosystems. The aims of this study were to determine how tree mortality influences the spatial patterns and interspecific associations of plant species, and what are the causes of tree mortality in a 1 hm² permanent plot in Baotianman National Nature Reserve, Nanyang City, Henan Province.
Methods We conducted field investigations in the plot and used spatial point pattern analysis to examine the spatial patterns and interspecific associations of 17 species prior to and following mortality.
Important findings (1) Most of the species in the study plot showed an aggregated distribution both pre- and post-mortality. However, the number of species showing aggregated distribution decreased and the number of species showing random distribution increased following the mortality event. (2) Most species were positively associated with Quercus aliena var. acuteserrata both pre- and post-mortality, while some had no apparent association. Following tree mortality, on fine scales, the number of species with positive associations increased, and the number of species with negative associations decreased. (3) Tree mortality was in consistency with the random death hypothesis. The interspecific associations of four species with Q. aliena var. acuteserrata completely changed following death. For most species, the spatial patterns and the interspecific association with Q. aliena var. acuteserrata either changed at minor scales or did not change. The variations in spatial patterns or interspecific associations were inconsistent among species. (4) The dead trees of Q. aliena var. acuteserrata were significantly associated with the living trees in 13 species, but the associations between dead and living trees were not in agreement with the changes in interspecific association following mortality. Only five living tree species competed with the dead trees of Q. aliena var. acuteserrata, and the competition between each of these species and Q. aliena var. acuteserrata intensified after individual death. Tree mortality is the result of a variety of factors. Although the tree mortality in the study plot was in accordance with the random death hypothesis, there were also a few individuals which were dead from competition.     

喀斯特常绿落叶阔叶林个体死亡对空间格局及种间关联性的影响

树木死亡作为森林生态系统中一个重要的生态学过程,研究其如何影响物种的空间分布格局及种间关联性,可以为揭示树木死亡机制及群落动态变化规律提供重要参考。本研究根据木论25 hm²喀斯特常绿落叶阔叶林动态监测样地两次木本植物调查数据,采用点格局分析方法,分析了树木死亡前后空间格局和种间关系的变化。结果表明: 样地内个体死亡前后主要表现为聚集分布,但死亡后聚集分布的物种比例较死亡前略微下降,小尺度上呈现随机分布的物种比例有所增加,个体的死亡呈非随机性死亡。在物种水平上,13个优势种的死树和活树之间在0～30 m各尺度上呈显著正关联,表明样地内优势物种个体之间种内和种间竞争作用不激烈;树木死亡前后种间关联性主要为正关联,且大部分物种种间关系在死亡前后并未发生变化,表明群落已发展到相对稳定的阶段;死亡后,在1～30 m尺度上表现为正关联的物种对比例有所增加,负关联和无关联的物种对在大部分尺度上有所减少,说明个体死亡发生后,各个物种之间的竞争压力在一定程度上有所缓解。     

Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone

Vegetation change from drought-induced mortality can alter ecosystem community structure, biodiversity, and services. Although drought-induced mortality of woody plants has increased globally with recent warming, influences of soil type, tree and shrub groups, and species are poorly understood. Following the severe 2002 drought in northern Arizona, we surveyed woody plant mortality and canopy dieback of live trees and shrubs at the forest–woodland ecotone on soils derived from three soil parent materials (cinder, flow basalt, sedimentary) that differed in texture and rockiness. Our first of three major findings was that soil parent material had little effect on mortality of both trees and shrubs, yet canopy dieback of trees was influenced by parent material; dieback was highest on the cinder for pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma). Ponderosa pine (Pinus ponderosa) dieback was not sensitive to parent material. Second, shrubs had similar mortality, but greater canopy dieback, than trees. Third, pinyon and ponderosa pines had greater mortality than juniper, yet juniper had greater dieback, reflecting different hydraulic characteristics among these tree species. Our results show that impacts of severe drought on woody plants differed among tree species and tree and shrub groups, and such impacts were widespread over different soils in the southwestern U.S. Increasing frequency of severe drought with climate warming will likely cause similar mortality to trees and shrubs over major soil types at the forest–woodland ecotone in this region, but due to greater mortality of other tree species, tree cover will shift from a mixture of species to dominance by junipers and shrubs. Surviving junipers and shrubs will also likely have diminished leaf area due to canopy dieback.     

Neighbour-regulated mortality: the influence of positive and negative density dependence on tree populations in species-rich tropical forests

Density‐dependent mortality has long been posited as a possible mechanism for the regulation of tropical forest tree density. Despite numerous experimental and phenomenological investigations, the extent to which such mechanisms operate in tropical forests remains unresolved because the demographical signature of density dependence has rarely been found in extensive investigations of established trees. This study used an individual‐based demographical approach to investigate the role of conspecific and heterospecific neighbourhood crowding on tree mortality in a Panamanian and a Malayan tropical forest. More than 80% of the species investigated at each site were found to exhibit density‐dependent mortality. Furthermore, most of these species showed patterns of mortality consistent with the Janzen–Connell hypothesis and the rarely explored hypothesis of species herd protection. This study presents some of the first evidence of species herd protection operating in tree communities.     

Growth and mortality patterns in a thinning canopy of post-hurricane regenerating rain forest in eastern Nicaragua (1990-2005)

One of the strongest hypothesis about the maintenance of tree species diversity in tropical areas is disturbance. In order to assess this, the effect of intensive natural disturbances on forest growth and mortality in a thinning canopy was studied after the landfall of hurricane Joan in 1988. We evaluated the growth and mortality rates of the 26 most common tree species of that forest in eastern Nicaragua. Permanent plots were established at two study sites within the damaged area. Growth and mortality rates of all individual trees > or = 3.18cm diameter at breast height were assessed annually from 1990 to 2005. During this period the forest underwent two phases: the building phase (marked by increased number of individuals of tree species present after the hurricane) and the canopy thinning phase (marked by increased competition and mortality). Our results from the thinning phase show that tree survival was independent of species identity and was positively related to the increase in growth rates. The analysis of mortality presented here aims to test the null hypothesis that individual trees die independently of their species identity. These findings were influenced by the mortality observed during the late thinning phase (2003-2005) and provide evidence in favor of a non-niche hypothesis at the thinning phase of forest regeneration.     

El Niño droughts and their effects on tree species composition and diversity in tropical rain forests

In this study I investigated the effects of the extreme, 1997/98 El Niño related drought on tree mortality and understorey light conditions of logged and unlogged tropical rain forest in the Indonesian province of East Kalimantan (Borneo). My objectives were to test (1) whether drought had a significant effect on tree mortality and understorey light conditions, (2) whether this effect was greater in logged than in undisturbed forest, (3) if the expected change in tree mortality and light conditions had an effect on Macaranga pioneer seedling and sapling densities, and (4) which (a)biotic factors influenced tree mortality during the drought. The 1997/1998 drought led to an additional tree mortality of 11.2, 18.1, and 22.7% in undisturbed, old logged and recently logged forest, respectively. Mortality was highest in logged forests, due to extremely high mortality of pioneer Macaranga trees (65.4%). Canopy openness was significantly higher during the drought than during the non-drought year (6.0, 8.6 and 10.4 vs 3.7, 3.8 and 3.7 in undisturbed, old logged and recently logged forest, respectively) and was positively correlated with the number of dead standing trees. The increase in light in the understorey was accompanied by a 30 to 300-fold increase in pioneer Macaranga seedling densities. Factors affecting tree mortality during drought were (1) tree species successional status, (2) tree size, and (3) tree location with respect to soil moisture. Tree density and basal area per surface unit had no influence on tree mortality during drought. The results of this study show that extreme droughts, such as those associated with El Niño events, can affect the tree species composition and diversity of tropical forests in two ways: (1) by disproportionate mortality of certain tree species groups and tree size classes, and (2) by changing the light environment in the forest understorey, thereby affecting the recruitment and growth conditions of small and immature trees.     

特大干旱对树木死亡的影响——以美国德克萨斯州东部森林为例

美国德克萨斯州在2011年经历了史上最严重的干旱,这一事件造成约3亿多株树木死亡。在大时空尺度上(面积约9×10~6 hm~2,时间跨度近20年),基于近1800个森林样地,4次周期性调查中的约209663株树木,使用主成份分析(PCA)和广义线性混合效应模型(GLM)回归,对树木死亡的时空差异及其干旱强度与长度对树木死亡造成的中长期复杂影响进行了研究。采用树木密度、树木基面积、林地年龄、样地调查时间间隔作为树木间的竞争指标,分析了造成大旱前后周期水平和年度水平上的样地树木死亡差异的原因。综合分析了不同地理区域、树木种组、胸径大小和林地起源的4个划分标准下树木死亡对死亡率的相对贡献。结果表明：松属树木的死亡率最低(7.92%);高度低、胸径小的树木的死亡率较大,分别为29.79%和26.00%。人工林的树木死亡率(10.26%)低于天然林(13.47%);西海湾平原生态区树木的死亡率在干旱后达到最大(22.27%);西南区的树木死亡率在干旱后也达到最大(13.78%);海拔和纬度对树木死亡率影响不明显。德州东部森林整体死亡格局形成原因较为复杂,各地理区域、林地起源、树木大小和不同树种,...     

Tornado damage of Quercus stellata and Quercus marilandica in the Cross Timbers,Oklahoma, USA

Questions: The Cross Timbers are a mosaic of savannas, grasslands and upland forests, occupying a significant portion of south‐central North America. Our questions here were (1) how does a severe tornado affect the two most dominant tree species of the area Quercus marilandica and Q. stellata with respect to damage and mortality; (2) how do such patterns vary as a function of tree size? What are the implications of disturbance for codominance in species‐poor systems? Location: The Cross Timbers in Oklahoma, USA. Methods: We established a 14.48‐ha permanent plot following a severe tornado in 2003. We identified, numbered and tagged each tree and recorded its diameter at breast height (DBH), spatial coordinates, status (dead or alive), and damage type. We examined (1) relative abundance before and after the tornado; (2) differences in damage and mortality, and (3) the influence of tree diameter on the probability of damage and mortality for each species. Results: Differences in species identity and tree characteristics were significantly related to tree mortality following the tornado, after accounting for spatial locations. The odds of mortality were 12.0 times greater for Q. marilandica than for Q. stellata. Such greater vulnerability of Q. marilandica versus Q. stellata was also reflected in changes in density and basal area. Tree diameter clearly influenced the damage and mortality pattern in Q. stellata; larger trees sustained more damage and mortality. However, Q. marilandica did not exhibit size‐dependent mortality. Conclusion: The tornado affected the two dominant species differently. The intra‐ and inter‐specific differences in windstorm susceptibility may allow coexistence of the two species and are potentially important in the dynamics of the Cross Timbers. Species more damaged might finally benefit from the wind disturbance due to their resprouting ability.     

Declining tree growth resilience mediates subsequent forest mortality in the US Mountain West

Climate change-triggered forest die-off is an increasing threat to global forests and carbon sequestration but remains extremely challenging to predict. Tree growth resilience metrics have been proposed as measurable proxies of tree susceptibility to mortality. However, it remains unclear whether tree growth resilience can improve predictions of stand-level mortality. Here, we use an extensive tree-ring dataset collected at ~3000 permanent forest inventory plots, spanning 13 dominant species across the US Mountain West, where forests have experienced strong drought and extensive die-off has been observed in the past two decades, to test the hypothesis that tree growth resilience to drought can explain and improve predictions of observed stand-level mortality. We found substantial increases in growth variability and temporal autocorrelation as well declining drought resistance and resilience for a number of species over the second half of the 20th century. Declining resilience and low tree growth were strongly associated with cross- and within-species patterns of mortality. Resilience metrics had similar explicative power compared to climate and stand structure, but the covariance structure among predictors implied that the effect of tree resilience on mortality could partially be explained by stand and climate variables. We conclude that tree growth resilience offers highly valuable insights on tree physiology by integrating the effect of stressors on forest mortality but may have only moderate potential to improve large-scale projections of forest die-off under climate change.     

Widespread drought‐induced tree mortality at dry range edges indicates that climate stress exceeds species' compensating mechanisms

Drought‐induced tree mortality is projected to increase due to climate change, which will have manifold ecological and societal impacts including the potential to weaken or reverse the terrestrial carbon sink. Predictions of tree mortality remain limited, in large part because within‐species variations in ecophysiology due to plasticity or adaptation and ecosystem adjustments could buffer mortality in dry locations. Here, we conduct a meta‐analysis of 50 studies spanning >100 woody plant species globally to quantify how populations within species vary in vulnerability to drought mortality and whether functional traits or climate mediate mortality patterns. We find that mortality predominantly occurs in drier populations and this pattern is more pronounced in species with xylem that can tolerate highly negative water potentials, typically considered to be an adaptive trait for dry regions, and species that experience higher variability in water stress. Our results indicate that climate stress has exceeded physiological and ecosystem‐level tolerance or compensating mechanisms by triggering extensive mortality at dry range edges and provides a foundation for future mortality projections in empirical distribution and mechanistic vegetation models.     

Slow lifelong growth predisposes Populus tremuloides trees to mortality

Widespread dieback of aspen forests, sometimes called sudden aspen decline, has been observed throughout much of western North America, with the highest mortality rates in the southwestern United States. Recent aspen mortality has been linked to drought stress and elevated temperatures characteristic of conditions expected under climate change, but the role of individual aspen tree growth patterns in contributing to recent tree mortality is less well known. We used tree-ring data to investigate the relationship between an individual aspen tree’s lifetime growth patterns and mortality. Surviving aspen trees had consistently higher average growth rates for at least 100 years than dead trees. Contrary to observations from late successional species, slow initial growth rates were not associated with a longer lifespan in aspen. Aspen trees that died had slower lifetime growth and slower growth at various stages of their lives than those that survived. Differences in average diameter growth between live and dead trees were significant (α = 0.05) across all time periods tested. Our best logistical model of aspen mortality indicates that younger aspen trees with lower recent growth rates and higher frequencies of abrupt growth declines had an increased risk of mortality. Our findings highlight the need for species-specific mortality functions in forest succession models. Size-dependent mortality functions suitable for late successional species may not be appropriate for species with different life history strategies. For some early successional species, like aspen, slow growth at various stages of the tree’s life is associated with increased mortality risk.     

Determinants of mortality across a tropical lowland rainforest community

Tree mortality is an important process determining forest dynamics. However, in species‐rich tropical forests it is largely unknown, how species differ in their response of mortality to resource availability and individual condition. We use a hierarchical Bayesian approach to quantify the impact of light availability, tree size and past growth on mortality of 284 woody species in a 50‐ha long‐term forest census plot in Barro Colorado Island, Panama. Light reaching each individual tree was estimated from yearly vertical censuses of vegetation density. Across the community, 78% of the species showed increasing mortality with increasing light. Considering species individually, just 29 species showed a significant response to light, all with increasing mortality at high light. Past growth had a significant impact on all but three species, with higher growth leading to lower mortality. For the majority of species, mortality decreased sharply with diameter in saplings, then levelled off or increased slightly towards the maximum diameter of the species. Diameter had the biggest impact on mortality, followed by past growth and finally light availability. Our results challenge many previous reports of higher mortality in shade, and we suggest that it is crucial to control for size effects when assessing the impact of environmental conditions on mortality.