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A tale of two species: climate-competition tradeoffs shape range limits according to tree life-history strategies
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Abstract
Adapting for competitiveness versus climatic stress tolerance constitutes a primary trade-off differentiating tree life-history strategies. This tradeoff likely influences where species’ range-limits occur, but such links are data-demanding to study and key mechanisms lack empirical support. Using an exceptionally rich dendroecological network, we assessed spatial variation in climate and competition effects on Picea abies and Fagus sylvatica throughout the Carpathian Ecoregion. Ring width synchrony aided in diagnosing how the prevalence of resource-limited (competition) and sink-limited (climate) growth changes with altitude and community composition. Contrasting growth patterns towards respective upper and lower range limits of Fagus and Picea reflected tradeoffs between competitive vs. cold-tolerant strategies. Fagus performance declined with altitudinal increases in climate sensitivity, but improved under interspecific competition. Picea growth increased towards the species’ lower range limit, but declined under interspecific competition. Warmer temperatures likely benefit competitively stronger species at mid elevations and thus imply range reductions for alpine conifers.
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Spruce-fir-beech mixed forests cover a large area in European mountain regions, with high ecological and socioeconomic importance. As elevation-zone systems they are highly affected by climate change, which is modifying species growth patterns and productivity shifts among species. The extent to which associated tree species can access resources and grow asynchronously may affect their resistance and persistence under climate change. Intra-specific synchrony in annual tree growth is a good indicator of species specific dependence on environmental conditions variability. However, little attention has been paid to explore the role of the inter-specific growth asynchrony in the adaptation of mixed forests to climate change. Here we used a database of 1790 tree-ring series collected from 28 experimental plots in spruce-fir-beech mixed forests across Europe to explore how spatio-temporal patterns of the intra-and inter-specific growth synchrony relate to climate variation during the past century. We further examined whether synchrony in growth response to inter-annual environmental fluctuations depended on site conditions. We found that the inter-specific growth synchrony was always lower than the intra-specific synchrony, for both high (inter-annual fluctuations) and low frequency (mid-to long-term)
Global change ecologists have often used trees under weak competition (e.g. dominant/codominant trees) to examine relationships between climatic change and tree growth. Scaling up these results to a forest relies on the assumption that the climatic change–tree growth relationship is not affected by tree‐level competition.
Using permanent sample plot data from the central Canadian boreal region where warming did not result in water deficit, we tested the above‐mentioned assumption by looking at whether the relationship between climatic change and tree growth varied with tree‐level competition, which was quantified using a modified Hegyi competition index.
We found that tree growth increased over time for trees under weak competition, but decreased for those under strong competition. The divergent temporal trends among trees under different levels of competition led to a non‐significant change in growth for our study plots. Growth increased with regional warming, atmospheric [CO2] and water availability for trees under weak competition, but not for those under strong competition.
Synthesis. Our results suggest that upscaling the growth responses of dominant/codominant trees to climate change to a forest or a region can lead to biased estimates. Tree‐level competition should be taken into account when expressing climatic change and tree growth relationships.
Species often respond to human‐caused climate change by shifting where they occur on the landscape. To anticipate these shifts, we need to understand the forces that determine where species currently occur. We tested whether a long‐hypothesised trade‐off between climate and competitive constraints explains where tree species grow on mountain slopes. Using tree rings, we reconstructed growth sensitivity to climate and competition in range centre and range margin tree populations in three climatically distinct regions. We found that climate often constrains growth at environmentally harsh elevational range boundaries, and that climatic and competitive constraints trade‐off at large spatial scales. However, there was less evidence that competition consistently constrained growth at benign elevational range boundaries; thus, local‐scale climate‐competition trade‐offs were infrequent. Our work underscores the difficulty of predicting local‐scale range dynamics, but suggests that the constraints on tree performance at a large‐scale (e.g. latitudinal) may be predicted from ecological theory.
Demographic events such as the birth and death of organisms are ubiquitous in nature. Disentangling the processes that underlie the demographic dynamics of species is fundamental to understanding how biodiversity is organized and will be re‐organized at multiple scales.
To provide new insights on multispecies demographic dynamics, I designed the present study to focus on local and biogeographic influences on mortality and recruitment rates for tree populations in forest biomes throughout the Japanese archipelago.
Most populations exhibited mortality that fell within the 95% confidence intervals of a neutral model (i.e. the expectation of demographic stochasticity). However, there were also important determinants that make population dynamics to deviate from the neutral expectation. Interspecific niche differentiation was important to reduce the mortality rate, regardless of the local rarity or ubiquity of a species. Although intraspecific aggregation that caused density‐dependent mortality was only significant for locally abundant species, the degree of isolation from a species’ central or optimal range determined mortality only for locally rare species. These differences between rare and common species provide important empirical quantification of aspects of population dynamics that have been less accounted for in ecology and biogeography studies.
Synthesis. Unifying different theories such as those rooted in local‐scale community ecology and macro‐scale biogeography is important. At this juncture, this study emphasizes that disentangling the interactive roles of demographic stochasticity and determinism that operate at different scales would be a clue to advance the field of ecology both in theory and practice.
A range of environmental factors regulate tree growth; however, climate is generally thought to most strongly influence year-to-year variability in growth. Numerous dendrochronological (tree-ring) studies have identified climate factors that influence year-to-year variability in growth for given tree species and location. However, traditional dendrochronology methods have limitations that prevent them from adequately assessing stand-level (as opposed to species-level) growth. We argue that stand-level growth analyses provide a more meaningful assessment of forest response to climate fluctuations, as well as the management options that may be employed to sustain forest productivity. Working in a mature, mixed-species stand at the Howland Research Forest of central Maine, USA, we used two alternatives to traditional dendrochronological analyses by (1) selecting trees for coring using a stratified (by size and species), random sampling method that ensures a representative sample of the stand, and (2) converting ring widths to biomass increments, which once summed, produced a representation of stand-level growth, while maintaining species identities or canopy position if needed. We then tested the relative influence of seasonal climate variables on year-to-year variability in the biomass increment using generalized least squares regression, while accounting for temporal autocorrelation. Our results indicate that stand-level growth responded most strongly to previous summer and current spring climate variables, resulting from a combination of individualistic climate responses occurring at the species- and canopy-position level. Our climate models were better fit to stand-level biomass increment than to species-level or canopy-position summaries. The relative growth responses (i.e. percent change) predicted from the most influential climate variables indicate stand-level growth varies less from to year-to-year than species-level or canopy-position growth responses. By assessing stand-level growth response to climate, we provide an alternative perspective on climate–growth relationships of forests, improving our understanding of forest growth dynamics under a fluctuating climate. This article is protected by copyright. All rights reserved.
Determining the drivers of shifting forest disturbance rates remains a pressing global change issue. Large-scale forest dynamics are commonly assumed to be climate driven, but appropriately scaled disturbance histories are rarely available to assess how disturbance legacies alter subsequent disturbance rates and the climate sensitivity of disturbance. We compiled multiple tree-ring based disturbance histories from primary Picea abies forest fragments distributed throughout five European landscapes spanning the Bohemian Forest and the Carpathian Mountains. The regional chronology includes 11 595 tree cores, with ring dates spanning the years 1750 to 2000, collected from 560 inventory plots in 37 stands distributed across a 1000 km geographic gradient, amounting to the largest disturbance chronology yet constructed in Europe. Decadal disturbance rates varied significantly through time and declined after 1920, resulting in widespread increases in canopy tree age. Approximately 75% of current canopy area recruited prior to 1900. Long-term disturbance patterns were compared to an historical drought reconstruction, and further linked to spatial variation in stand structure and contemporary disturbance patterns derived from LANDSAT imagery. Historically, decadal Palmer drought severity index minima corresponded with higher rates of canopy removal. The severity of contemporary disturbances increased with each stand's estimated time since last major disturbance, increased with mean diameter and declined with increasing within-stand structural variability. Reconstructed spatial patterns suggest that high small-scale structural variability has historically acted to reduce large-scale susceptibility and climate sensitivity of disturbance. Reduced disturbance rates since 1920, a potential legacy of high 19th century disturbance rates, have contributed to a recent region-wide increase in disturbance susceptibility. Increasingly common high-severity disturbances throughout primary Picea forests of Central Europe should be reinterpreted in light of both legacy effects (resulting in increased susceptibility) and climate change (resulting in increased exposure to extreme events). This article is protected by copyright. All rights reserved.
The role of density dependence in shaping spatial patterns in tree distributions presumably changes throughout stand development. However, empirical investigations into developmental processes are often limited by a lack of long-term data on disturbance history, which further limits the ability to assess the role of spatial variation in site conditions (e.g., slope, aspect, mean annual temperature). This study included data from 289 plots within 26 primary forest stands of the Carpathian Mountains; stands were dominated by Norway spruce (Picea abies) and driven by mixed-severity disturbance regimes. We assessed spatial patterns in living tree positions, tree diameters, and the relative position of living trees to dead trees. Random forest classification was used to discriminate between disturbance history, tree density, and site conditions and their effects on the observed spatial patterns. At the stand scale, distances between trees of equal diameter were more uniform that expected (tree diameter was showing repulsion), while tree positions and dead trees were mostly distributed randomly. The processes that best explained the spatial patterns were identified as self-thinning mortality and past disturbances (100–150 yr). This study demonstrated that the plot and stand-scale spatial patterns resulted from the combination of past disturbances and density-dependent legacies derived from earlier forest development stages.
Significance
Forests dominate carbon fluxes in terrestrial ecosystems. We demonstrate how an intensified climatic influence on tree growth during the last 120 y has increased spatial synchrony in annual ring-width patterns within contrasting (boreal and Mediterranean) Eurasian biomes and on broad spatial scales. Current trends in tree growth synchrony are related to regional changes in climate factors controlling productivity, overriding local and taxonomic imprints on forest carbon dynamics. Enhanced synchrony is becoming a widespread, although regionally dependent, phenomenon related to warmer springs and increased temperature variability in high latitudes and to warmer winters and drier growing seasons in mid-latitudes.
Duncan, R.P. (1989). An evaluation of errors in tree age estimates based on increment cores in kahikatea (Dacrycarpus dacrydioides). New Zealand Natural Sciences 16: 31-37. Twelve kahikatea (Dacrycarpus dacrydioides) discs were used to assess the likely errors associated with estimating tree age from growth ring counts in increment cores. Two major sources of error were examined: (1) Failure of the increment core to pass through the tree's chronological centre. A geometric model is developed for estimating the distance to the chronological centre in cores where the arcs of the inner rings are visible. The mean percentage error from 84 cores that passed within 50 mm of the chronological centre was ± 35% corresponding to a mean absolute error of ± 21 years. The majority of this error is due to growth rate differences between the missing radius and the measured part of the core. (2) Missing rings. The average age underestimate from 48 cores due to missing rings was 13%. A significant correlation between radius length and age under estimate (r = 0.81) suggests that sampling along the longest radii will reduce this error. The average age underestimate due to missing rings from cores located along the longest radii of the twelve samples was 3%.
The Carpathians are the longest mountain range in Europe and a geographic barrier between Central Europe, Eastern Europe, and the Balkans. To investigate the climate of the area, the CARPATCLIM project members collected, quality-checked, homogenized, harmonized, and interpolated daily data for 16 meteorological variables and many derived indicators related to the period 1961–2010. The principal outcome of the project is the Climate Atlas of the Carpathian Region, hosted on a dedicated website (www.carpatclim-eu.org) and made of high-resolution daily grids (0.1° × 0.1°) of all variables and indicators at different time steps. In this article, we analyze the spatial and temporal variability of 10 variables: minimum, mean, and maximum temperature, daily temperature range, precipitation, cloud cover, relative sunshine duration, relative humidity, surface air pressure, and wind speed at 2 m. For each variable, we present the gridded climatologies for the period 1961–2010 and discuss the linear trends both on an annual and seasonal basis. Temperature was found to increase in every season, in particular in the last three decades, confirming the trends occurring in Europe; wind speed decreased in every season; cloud cover and relative humidity decreased in spring, summer, and winter, and increased in autumn, while relative sunshine duration behaved in the opposite way; precipitation and surface air pressure showed no significant trend, though they increased slightly on an annual basis. We also discuss the correlation between the variables and we highlight that in the Carpathian Region positive and negative sunshine duration anomalies are highly correlated to the corresponding temperature anomalies during the global dimming (1960s and 1970s) and brightening (1990s and 2000s) periods.
Questions: How have the historical frequency and severity of natural disturbances
in primary Picea abies forests varied at the forest stand and landscape level
during recent centuries? Is there a relationship between physiographic attributes
and historical patterns of disturbance severity in this system?
Location: Primary P. abies forests of the Eastern Carpathian Mountains, Romania;
a region thought to hold the largest concentration of primary P. abies forests
in Europe’s temperate zone.
Methods: We used dendrochronological methods applied to many plots over a
large area (132 plots representing six stands in two landscapes), thereby providing
information at both stand and landscape levels. Evidence of past canopy disturbance
was derived from two patterns of radial growth: (1) abrupt, sustained
increases in growth (releases) and (2) rapid early growth rates (gap recruitment).
Thesemethods were augmented with non-metricmultidimensional scaling
to facilitate the interpretation of factors influencing past disturbance.
Results: Of the two growth pattern criteria used to assess past disturbance, gap
recruitment was the most common, representing 80% of disturbance evidence
overall. Disturbance severities varied over the landscape, including stand-replacing
events, as well as low- and intermediate-severity disturbances. More than
half of the study plots experienced extreme-severity disturbances at the plot
level, although they were not always synchronized across stands and landscapes.
Plots indicating high-severity disturbances were often spatially clustered
(indicating disturbances up to 20 ha), while this tendency was less clear for lowand
moderate-severity disturbances. Physiographic attributes such as altitude
and land form were only weakly correlated with disturbance severity. Historical
documents suggest windstorms as the primary disturbance agent, while the role
of bark beetles (Ips typographus) remains unclear.
Conclusions: The historical disturbance regime revealed in this multi-scale
study is characterized by considerable spatial and temporal heterogeneity,which
could be seen among plots within stands, among stands within landscapes and
between the two landscapes. When the disturbance regime was evaluated at
these larger scales, the entire range of disturbance severity was revealed within
this landscape.
Winter physiology of woody plants is a key issue in temperate biomes. Here, we investigated different frost resistance mechanisms on 1-year-old branches of 11 European tree species from November until budburst: (i) frost hardiness of living cells (by electrolyte leakage method), (ii) winter embolism sensitivity (by percentage loss of conductivity: PLC) and (iii) phenological variation of budburst (by thermal time to budburst). These ecophysiological traits were analyzed according to the potential altitudinal limit, which is highly related to frost exposure. Seasonal frost hardiness and PLC changes are relatively different across species. Maximal PLC observed in winter (PLCMax) was the factor most closely related to potential altitudinal limit. Moreover, PLCMax was related to the mean hydraulic diameter of vessels (indicating embolism sensitivity) and to osmotic compounds (indicating ability of living cells to refill xylem conducting elements). Winter embolism formation seems to be counterbalanced by active refilling from living cells. These results enabled us to model potential altitudinal limit according to three of the physiological/anatomical parameters studied. Monitoring different frost resistance strategies brings new insights to our understanding of the altitudinal limits of trees.
Angiosperm radiation in the Cretaceous is thought to have profoundly diminished the success of the conifers, the other major woody plant group present at the time. However, today the conifers persist and often thrive despite their supposed inferiority in vegetative and reproductive function. By exploring this apparent conflict for global tree dominance, we seek here to reveal patterns that explain not only how the allegedly inferior conifers persist among angiosperms but also why some conifer groups became extinct in the Cretaceous. We find that despite the profound contrast between the dominant conifer families in the Southern and Northern Hemispheres, all conifers can be characterized by a common set of functional attributes that allow them to exist in an important group of niches, from high latitudes to the equator. In these environments, conifers are often highly efficient at outcompeting, outliving, or outsurviving angiosperms. Hence, we conclude that conifer success cannot be dismissed as being uniquely associated with habitats that are unfavorable for angiosperms.
European forests, covering more than 2 mio km2 or 32% of the land surface1, are to a large extent intensively managed and support an important timber industry. Climate change is expected to strongly effect tree species distribution within these forests2,3. Climate and land use are currently undergoing rapid changes4, with initial range shifts already visible5. However, discussions on the consequences of biome shifts have concentrated on ecological issues6. Here we show that forecasted changes in temperature and precipitation may have severe economic consequences. Based on our model results, the expected value of European forestland will decrease due to the decline of economically valuable species in the absence of effective countermeasures. We found that by 2100 – depending on interest rate (ir) and climate scenario applied – this loss varies between 14 and 50% (mean: 28% for ir=2%) of the current value of forestland in Europe, excluding Russia, and may total several hundred billion Euro. Our model shows that – depending on different realizations of three climate scenarios – by 2100 between 21 and 60% (mean: 34%) of European forestlands will only be suitable for a Mediterranean oak forest type with low economic returns for forest owners and the timber industry and reduced carbon sequestration.
This paper provides an overview of climatic changes that have been observed during the past century at certain high-elevation sites, and changes in a more distant past documented by a variety of climate-sensitive environmental indicators, such as tree-rings and alpine glaciers, that serve as a measure of the natural variability of climate in mountains over longer time scales.
Detailed studies such as those found in this special issue of Climatic Change, as well as those noted in this review, for the mountain regions of the world, advance our understanding in a variety of ways. They are not only helpful to characterize present and past climatological features in the mountainous zones, but they also provide useful information to the climate modeling community. Because of the expected refinements in the physical parameterization of climate models in coming years, and the probable increase in the spatial resolution of GCMs, the use of appropriate data from high elevation sites will become of increasing importance for model initialization, verification, and intercomparison purposes. The necessity of accurate projections of climate change is paramount to assessing the likely impacts of climate change on mountain biodiversity, hydrology and cryosphere, and on the numerous economic activities which take place in these regions.
We have developed extensions of traditional distance-dependent, spatial competition analyses that estimate the magnitude of the competitive effects of neighboring trees on target tree growth as a function of the species, size, and distance to neighboring trees. Our analyses also estimate inter- and intra-specific competition coefficients and explicitly partition the competitive effects of neighbors into the effects of shading versus crowding. We tested the method using data from forests of northern, interior British Columbia dominated by western hemlock (Tsuga heterophylla (Raf.) Sarg.) and western redcedar (Thuja plicata Donn ex D. Don). For both species, the most parsimonious regression models included terms for the effects of tree size, crowding, and shading and separate competitive effects of four different groups of competing species. The models explained 33%-59% of the variation in radial growth of the two species. For both species, growth declined more steeply as a function of crowding than shading. There was striking asymmetry in the strength of interspecific competition between hemlock and redcedar, with crowding by hemlock having a strong per capita effect on redcedar, while crowding by redcedar had relatively little effect on the radial growth of hemlock.
Variation in plant functional traits results from evolutionary and environmental drivers that operate at a variety of different scales, which makes it a challenge to differentiate among them. In this article we describe patterns of functional trait variation and trait correlations within and among habitats in relation to several environmental and trade-off axes. We then ask whether such patterns reflect natural selection and can be considered plant strategies. In so doing we highlight evidence that demonstrates that
(1) patterns of trait variation across resource and environmental gradients (light, water, nutrients, and temperature) probably reflect adaptation, (2) plant trait variation typically involves multiple-correlated traits that arise because of inevitable trade-offs among traits and across levels of whole-plant integration and that must be understood from a wholeplant perspective, and
(3) such adaptation may be globally generalizable for like conditions; i.e., the set of traits (collections of traits in syndromes) of taxa can be considered as “plant strategies.”
Does climate determine species' ranges? Rapid rates of anthropogenic warming make this classic ecological question especially relevant. We ask whether climate controls range limits by quantifying relationships between climatic variables (precipitation, temperature) and tree growth across the altitudinal ranges of six Pacific Northwestern conifers on Mt. Rainier, Washington, USA. Results for three species (Abies amabilis, Callitropsis nootkatensis, Tsuga mertensiana) whose upper limits occur at treeline (> 1600 m) imply climatic controls on upper range limits, with low growth in cold and high snowpack years. Annual growth was synchronized among individuals at upper limits for these high-elevation species, further suggesting that stand-level effects such as climate constrain growth more strongly than local processes. By contrast, at lower limits climatic effects on growth were weak for these high-elevation species. Growth-climate relationships for three low-elevation species (Pseudotsuga menziesii, Thuja plicata, Tsuga heterophylla) were not consistent with expectations of climatic controls on upper limits, which are located within closed-canopy forest (< 1200 m). Annual growth of these species was poorly synchronized among individuals. Our results suggest that climate controls altitudinal range limits at treeline, while local drivers (perhaps biotic interactions) influence growth in closed-canopy forests. Climate-change-induced range shifts in closed-canopy forests will therefore be difficult to predict accurately.
• Existing growth and yield plots of pure and mixed stands of Norway spruce (Picea abies (L.) H. Karst.) and European beech (Fagus sylvatica L.) were aggregated in order to unify the somewhat scattered sources of information currently available, as well as to develop a sound working hypothesis about mixing effects. The database contains information from 23 long-term plots, covering an ecological gradient from nutrient poor and dry to nutrient rich and moist sites throughout Central Europe.
• An empirically formed interaction model showed, that depending on the site conditions, dry mass growth in mixed stands can range from −46% to +138 % of the growth yielded by a scaled combination of pure stands at equal mixing proportions.
• Drawing from the interaction model, overyielding of the mixed stands appears to be triggered by two separate mechanisms. On poor sites, where significant overyielding is commonly found, facilitation by beech offsets nutrient-related growth limitations in spruce. In contrast, overyielding of mixed stands occurs less frequently on rich sites, and appears to be based on an admixture effect, with spruce reducing the severe intra-specific competition common in pure beech stands.
• It was concluded that silviculture can accelerate growth of spruce by beech admixtures on poor sites, while growth of beech can be promoted by admixture of spruce, particularly on excellent sites.
Hydraulic conductivity in the terminal branches of mature beech trees (Fagus sylvatica L.) decreased progressively during winter and recovered in the spring. The objective of this study was to determine the mechanisms
involved in recovery. Two periods of recovery were identified. The first recovery of hydraulic conductivity occurred early
in the spring, before bud break, and was correlated with the occurrence of positive xylem pressure at the base of the tree
trunk. Active refilling of the embolized vessels caused the recovery. The second recovery of hydraulic conductivity occurred
after bud break and was correlated with the onset of cambial activity. Formation of new functional vessels, leading to an
increase in xylem diameter, was largely responsible for the increase in xylem conductivity. The two mechanisms were complementary:
active refilling of embolized vessels occurred mostly in the root and the trunk, whereas formation of new functional vessels
occurred mainly in young terminal shoots.
Shifting forest dynamics
Forest dynamics are the processes of recruitment, growth, death, and turnover of the constituent tree species of the forest community. These processes are driven by disturbances both natural and anthropogenic. McDowell et al. review recent progress in understanding the drivers of forest dynamics and how these are interacting and changing in the context of global climate change. The authors show that shifts in forest dynamics are already occurring, and the emerging pattern is that global forests are tending toward younger stands with faster turnover as old-growth forest with stable dynamics are dwindling.
Science , this issue p. eaaz9463
While shifting disturbance rates and climate change have major implications for the structure of contemporary forests through their effects on adult tree mortality, the responses of regenerating trees to disturbances and environmental variation will ultimately determine the structure and functioning of forests in the future. Assessing the resilience of forests to changing conditions requires information on what constrains tree performance during recruitment and whether recruitment dynamics have changed throughout history.
We analyzed growth patterns in a large sample of tree cores (n = 14 793) collected from primary Picea forests throughout the Carpathian Mountains. Growth rate anomalies recorded in tree-rings permitted the reconstruction of several key recruitment and disturbance parameters: (1) whether individuals were recruited after a period of competitive suppression (Released Trees; RT; 66% of trees) or immediately following gap formation (Gap Recruited Trees; GRT; 33%), (2) growth rates during recruitment, (3) the duration of recruitment and (4) historical disturbance severity variation.
High neighborhood density led to lower growth rates in RTs, but favored a higher growth rate in GRTs. Winter temperatures were positively correlated with Picea growth during recruitment, GRTs were also more sensitive to winter precipitation. Recent increases in growth during recruitment and reductions in recruitment intervals suggest that rates of canopy replacement have increased over recent decades.
Assessments of forest resilience must recognize that constraints on tree growth differ during recruitment and interact with disturbance severity. An individual's experience prior to competitive release and factors altering the immediate abiotic conditions of a recruiting individual (competition and disturbance severity) are important determinants of canopy replacement rates; these recruitment parameters will certainly interact with shifting disturbance regimes. Ultimately, increasing growth rates and decreasing recruitment intervals suggest that forest dynamics are accelerating, and are potentially compensating for recent increases in tree mortality rates.
Climatic constraints on tree growth mediate an important link between terrestrial and atmospheric carbon pools. Tree rings provide valuable information on climate‐driven growth patterns, but existing data tend to be biased towards older trees on climatically extreme sites. Understanding climate change responses of biogeographic regions requires data that integrate spatial variability in growing conditions and forest structure. We analyzed both temporal (c. 1901‐2010) and spatial variation in radial growth patterns in 9 876 trees from fragments of primary Picea abies forests spanning the latitudinal and altitudinal extent of the Carpathian arc. Growth was positively correlated with summer temperatures and spring moisture availability throughout the entire region. However, important seasonal variation in climate responses occurred along geospatial gradients. At northern sites, winter precipitation and October temperatures of the year preceding ring formation were positively correlated with ring width. In contrast, trees at the southern extent of the Carpathians responded negatively to warm and dry conditions in autumn of the year preceding ring formation. An assessment of regional synchronization in radial growth variability showed temporal fluctuations throughout the 20th century linked to the onset of moisture limitation in southern landscapes. Since the beginning of the study period, differences between high and low elevations in the temperature sensitivity of tree growth generally declined, while moisture sensitivity increased at lower elevations. Growth trend analyses demonstrated changes in absolute tree growth rates linked to climatic change, with basal area increments in northern landscapes and lower altitudes responding positively to recent warming. Tree growth has predominantly increased with rising temperatures in the Carpathians, accompanied by early indicators that portions of the mountain range are transitioning from temperature to moisture limitation. Continued warming will alleviate large‐scale temperature constraints on tree growth, giving increasing weight to local drivers that are more challenging to predict. This article is protected by copyright. All rights reserved.
The demand for large-scale and long-term information on tree growth is increasing rapidly as environmental change research strives to quantify and forecast the impacts of continued warming on forest ecosystems. This demand, combined with the now quasi-global availability of tree-ring observations, has inspired researchers to compile large tree-ring networks to address continental or even global-scale research questions. However, these emergent spatial objectives contrast with paleo-oriented research ideas that have guided the development of many existing records. A series of challenges related to how, where, and when samples have been collected is complicating the transition of tree rings from a local to a global resource on the question of tree growth. Herein, we review possibilities to scale tree-ring data (A) from the sample to the whole tree, (B) from the tree to the site, and (C) from the site to larger spatial domains. Representative tree-ring sampling supported by creative statistical approaches is thereby key to robustly capture the heterogeneity of climate-growth responses across forested landscapes. We highlight the benefits of combining the temporal information embedded in tree rings with the spatial information offered by forest inventories and earth observations to quantify tree growth and its drivers. In addition, we show how the continued development of mechanistic tree-ring models can help address some of the non-linearities and feedbacks that complicate making inference from tree-ring data. By embracing scaling issues, the discipline of dendrochronology will greatly increase its contributions to assessing climate impacts on forests and support the development of adaptation strategies.
https://authors.elsevier.com/c/1XWsP-4PRq7xR (free access until Sep 27)
Primary forests are characterized by high vertical and horizontal stand diversity, which provides habitat for a diverse range of species with complex habitat requirements. Detailed knowledge of related ecological processes and habitat development of primary forest species are essential to inform forest management and biodiversity conservation decisions, but relationships are not well documented. We collected dendrochronological data and inventoried numerous structural elements in permanent plots throughout the primary temperate forests within the Carpathian Mountains. We fit and compared multiple predictive models to quantify the importance of 200 years of natural disturbance dynamics on the occurrence probability of an umbrella species – the capercaillie (Tetrao urogallus). We showed that a mixed-severity disturbance regime ranging from low through moderate to high severity disturbances is required to generate diverse forest habitats suitable for capercaillie. The variation in natural disturbance severity and its timing promoted key structural habitat elements, such as low natural regeneration density, low mature tree density, high ground vegetation cover, availability of forest gaps, and abundance of standing deadwood. This study demonstrates the importance of natural disturbance in maintaining the variety of conditions necessary to support primary forest specialist species. Managers of protected areas should be mindful that natural disturbances generate habitat for the capercaillie in mountain Norway spruce forests. Further intervention is unnecessary. Conservation planning and forest reserve design should shift focus to the large-scale spatial requirements needed to ensure that a wide range of forest developmental phases are represented in protected areas.
Knowledge of the functional consequences of biodiversity is increasing through studies of both experimental systems and natural systems. Community assembly theory has also helped to reveal the causes of biodiversity organization. However, the causes and consequences of biodiversity have been discussed in parallel and simultaneous consideration of both has been limited, even though they are both influenced by regional environmental conditions.
To understand the relationship between biodiversity and productivity, I focused on the linkages between the number of tree species and biomass productivity across forest biomes under a range of bioclimatic conditions.
I found that high tree diversity generally increased productivity. This was primarily due to a selection process that results from a high probability of having high‐performance species and their dominance at high diversity, regardless of the biome. In less‐productive biomes, the residual importance of diversity, which likely reflects other forms of biological interactions (including species complementarity), increased productivity.
These findings of differential diversity effects under different environmental conditions are consistent with the existing theory of community assembly, which predicts a shift in the assembly process from stochasticity to determinism with increasing environmental harshness. Analyses based on functional trait diversity also supported this theory: stochastic assembly (resulting in the selection effect) and deterministic assembly (possibly resulting from interspecific niche differences) became more important in productive and less‐productive biomes, respectively.
Synthesis . Increasing our understanding of the causalities between diversity and other characteristics, such as productivity, is crucial, particularly for forest ecosystems, because of the increasing interest in productivity‐related ecosystem services supported by diverse assemblages of trees.
Climate is widely assumed to influence physiological and demographic processes in trees, and hence forest composition, biomass and range limits. Growth in trees is an important barometer of climate change impacts on forests as growth is highly correlated with other demographic processes including tree mortality and fecundity.
We investigated the main drivers of diameter growth for five common tree species occurring in the Rocky Mountains of the western United States using nonlinear regression methods. We quantified growth at the individual tree level from tree core samples collected across broad environmental gradients. We estimated the effects of both climate variation and biotic interactions on growth processes and tested for evidence that disjunct populations of a species respond differentially to climate.
Relationships between tree growth and climate varied by species and location. Growth in all species responded positively to increases in annual moisture up to a threshold level. Modest linear responses to temperature, both positive and negative, were observed at many sites. However, model results also revealed evidence for differentiated responses to local site conditions in all species. In severe environments in particular, growth responses varied nonlinearly with temperature. For example, in northerly cold locations pronounced positive growth responses to increasing temperatures were observed. In warmer southerly climates, growth responses were unimodal, declining markedly above a threshold temperature level.
Net effects from biotic interactions on diameter growth were negative for all study species. Evidence for facilitative effects was not detected. For some species, competitive effects more strongly influenced growth performance than climate. Competitive interactions also modified growth responses to climate to some degree.
Synthesis . These analyses suggest that climate change will have complex, species‐specific effects on tree growth in the Rocky Mountains due to nonlinear responses to climate, differentiated growth processes that vary by location and complex species interactions that impact growth and potentially modify responses to climate. Thus, robust model simulations of future growth responses to climate trends may need to integrate realistic scenarios of neighbourhood effects as well as variability in tree performance attributed to differentiated populations.
Climate and competition are often presented from two opposing views of the dominant driver of individual tree growth and species distribution in temperate forests, such as those in the eastern United States. Previous studies have provided abundant evidence indicating that both factors influence tree growth, and we argue that these effects are not independent of one another and rather that interactions between climate, competition, and size best describe tree growth. To illustrate this point, we describe the growth responses of five common eastern tree species to interacting effects of temperature, precipitation, competition, and individual size using maximum likelihood estimation. Models that explicitly include interactions among these four factors explained over half of the variance in annual growth for four out of five species using annual climate. Expanding temperature and precipitation analyses to include seasonal interactions resulted in slightly improved models with a mean R-2 of 0.61 (SD 0.10). Growth responses to individual factors as well their interactions varied greatly among species. For example, growth sensitivity to temperature for Quercus rubra increased with maximum annual precipitation, but other species showed no change in sensitivity or slightly reduced annual growth. Our results also indicate that three-way interactions among individual stem size, competition, and temperature may determine which of the five co-occurring species in our study could have the highest growth rate in a given year. Continued consideration and quantification of interactions among climate, competition, and individual-based characteristics are likely to increase understanding of key biological processes such as tree growth. Greater parameterization of interactions between traditionally segregated factors such as climate and competition may also help build a framework to reconcile drivers of individual-based processes such as growth with larger-scale patterns of species distribution.
A long-standing theory, originating with Darwin, suggests that abiotic forces set species range limits at high latitude, high elevation, and other abiotically 'stressful' areas, while species interactions set range limits in apparently more benign regions. This theory is of considerable importance for both basic and applied ecology, and while it is often assumed to be a ubiquitous pattern, it has not been clearly defined or broadly tested. We review tests of this idea and dissect how the strength of species interactions must vary across stress gradients to generate the predicted pattern. We conclude by suggesting approaches to better test this theory, which will deepen our understanding of the forces that determine species ranges and govern responses to climate change.
Multiple hypotheses have been put forward to explain the rise of angiosperms to ecological dominance following the Cretaceous. A unified scheme incorporating all these theories appears to be an inextricable knot of relationships, processes and plant traits. Here, we revisit these hypotheses, categorising them within frameworks based on plant carbon economy, resistance to climatic stresses, nutrient economy, biotic interactions and diversification. We maintain that the enigma remains unresolved partly because our current state of knowledge is a result of the fragmentary nature of palaeodata. This lack of palaeodata limits our ability to draw firm conclusions. Nonetheless, based on consistent results, some inferences may be drawn. Our results indicate that a complex multidriver hypothesis may be more suitable than any single-driver theory. We contend that plant carbon economy and diversification may have played an important role during the early stages of gymnosperms replacement by angiosperms in fertile tropical sites. Plant tolerance to climatic stresses, plant nutrition, biotic interactions and diversification may have played a role in later stages of angiosperm expansion within temperate and harsh environments. The angiosperm knot remains partly tied, but to unravel it entirely will only be feasible if new discoveries are made by scientific communities.
This review reports on the processes associated with carbon transfer and metabolism in leaves and growing organs and the role of long‐distance transport and vascular links in the regulation of carbon partitioning in plants. Partitioning is clearly influenced by both the supply and demand for photosynthate and is moderated by vascular connections and the storage capacity of the leaves and pathway tissues. However there appears to be little more than circumstantial evidence either that short distance transfer of carbon within either the source or the sink, or that long‐distance transport in the phloem, are limiting photosynthesis or growth directly. Although individual biochemical and physiological processes relating to photosynthesis and growth may be well understood, the factors primarily responsible for the control of carbon partitioning in plants have not been clearly identified. There is a need for a greater understanding of organ initiation and development (source and sink formation and potential size), the clear identification of whether growth is sink or source limited (including possible sink‐controlled photosynthesis) and a detailed assessment of the role of storage in buffering developmental and environmental changes in sink and source activity. Also more information is needed on the role of hormonal and nutritional factors in regulating source and sink activity (organ interactions not directly associated with carbon transfer).
CONTENTS
Summary
341
I.
Introduction
342
II.
General source‐sink relationships
342
III.
Control at the source
345
IV.
The utilization of photosynthate: sink characteristics and limitations
353
V.
Vascular constraints and temporary storage
360
VI.
Concluding comments
366
Acknowledgements
366
References
367
There is a need for a consistent forest restoration strategy for the `Black Triangle', Central Europe. In the past 50 years, forests in this area have been heavily affected by industrial pollution. Recently, the amount of pollutants has decreased. This means that effective forest restoration programmes can be started. Forest decline must be seen as an ecological disturbance which cannot be solved by applying technical measures only. An ecosystem approach to forest restoration must be introduced into restoration policy and management. Basic principles and working methods of such an approach are briefly described and proposals are made for forest restoration policy, management and research in the area.
The use of both linear and generalized linear mixed‐effects models ( LMM s and GLMM s) has become popular not only in social and medical sciences, but also in biological sciences, especially in the field of ecology and evolution. Information criteria, such as Akaike Information Criterion ( AIC ), are usually presented as model comparison tools for mixed‐effects models.
The presentation of ‘variance explained’ ( R ² ) as a relevant summarizing statistic of mixed‐effects models, however, is rare, even though R ² is routinely reported for linear models ( LM s) and also generalized linear models ( GLM s). R ² has the extremely useful property of providing an absolute value for the goodness‐of‐fit of a model, which cannot be given by the information criteria. As a summary statistic that describes the amount of variance explained, R ² can also be a quantity of biological interest.
One reason for the under‐appreciation of R ² for mixed‐effects models lies in the fact that R ² can be defined in a number of ways. Furthermore, most definitions of R ² for mixed‐effects have theoretical problems (e.g. decreased or negative R ² values in larger models) and/or their use is hindered by practical difficulties (e.g. implementation).
Here, we make a case for the importance of reporting R ² for mixed‐effects models. We first provide the common definitions of R ² for LM s and GLM s and discuss the key problems associated with calculating R ² for mixed‐effects models. We then recommend a general and simple method for calculating two types of R ² (marginal and conditional R ² ) for both LMM s and GLMM s, which are less susceptible to common problems.
This method is illustrated by examples and can be widely employed by researchers in any fields of research, regardless of software packages used for fitting mixed‐effects models. The proposed method has the potential to facilitate the presentation of R ² for a wide range of circumstances.
A number of parallels are shown between economic theory and plant resource usage. An outline of economic theory leads to 5 predictions concerning plant processes. Resource acquisition, storage, growth, resource loss and sexual reproduction are considered in the light of these predictions. Plants adjust phenology and life history patterns to acquire resources when they are cheap, store these internally and utilise them when conditions are favourable for growth. Plants continue to produce leaves (and perhaps roots) only until the marginal revenue from this increased production is equal to the marginal cost. Plants adjust allocation so that their limitation of growth is more nearly equal for all resources. Plants adjust physiologically to changes in resource availability to reduce extreme exchange ratios; the balance of internal reserves within the plant thereby approaches the proportions that are optimal for growth of most plants.-P.J.Jarvis
Gymnosperms, and conifers in particular, are sometimes very productive trees yet angiosperms dominate most temperate and tropical vegetation. Current explanations for angiosperm success emphasize the advantages of insect pollination and seed dispersal by animals for the colonization of isolated habitats. Differences between gymnosperm and angiosperm reproductive and vegetative growth rates have been largely ignored. Gymnosperms are all woody, perennial and usually have long reproductive cycles. Their leaves are not as fully vascularized as those of angiosperms and are more stereotyped in shape and size. Gymnosperm tracheids are generally more resistant to solute flow than angiosperm vessels. A consequence of the less efficient transport system is that maximum growth rates of gymnosperms are lower than maximum growth rates of angiosperms in well lit, well watered habitats. Gymnosperm seedlings may be particularly uncompetitive since their growth depends on a single cohort of relatively inefficient leaves. Later, some gymnosperms attain a higher productivity than co-occurring angiosperm trees by accumulating several cohorts of leaves with a higher total leaf area.
The ongoing enrichment of the atmosphere with CO 2 raises the question of whether growth of forest trees, which represent close to 90% of the global biomass carbon, is still carbon limited at current concentrations of close to 370 p.p.m. As photosynthesis of C3 plants is not CO 2 ‐saturated at such concentrations, enhanced ‘source activity’ of leaves could stimulate ‘sink activity’ (i.e. growth) of plants, provided other resources and developmental controls permit. I explore current levels of non‐structural carbon in trees in natural forests in order to estimate the potential for a carbon‐driven stimulation of growth.
The concentration of non‐structural carbohydrates (NSC) in tree tissues is considered a measure of carbon shortage or surplus for growth. A periodic reduction of NSC pools indicates either that carbon demand exceeds con‐current supply, or that both source and sink activity are low. A steady, very high NSC concentration is likely to indicate that photosynthesis fully meets, or even exeeds, that needed for growth (surplus assimilates accumulate).
The analysis presented here considers data for mature trees in four climatic zones: the high elevation treeline (in Mexico, the Alps and Northern Sweden), a temperate lowland forest of central Europe, Mediterranean sclerophyllous woodland and a semideciduous tropical forest in Panama.
In all four climatic regions, periods of reduced or zero growth show maximum C‐loading of trees (source activity exceeding demand), except for dry midsummer in the Mediterranean. NSC pools are generally high throughout the year, and are not significantly affected by mass fruiting episodes.
It is concluded that, irrespective of the reason for its periodic cessation, growth does not seem to be limited by carbon supply. Instead, in all the cases examined, sink activity and its direct control by the environment or developmental constraints, restricts biomass production of trees under current ambient CO 2 concentrations.
The current carbohydrate charging of mature wild trees from the tropics to the cold limit of tree growth suggests that little (if any) leeway exists for further CO 2 ‐fertilization effects on growth.
The expected upward shift of trees due to climate warming is supposed to be a major threat to range-restricted high-altitude species by shrinking the area of their suitable habitats. Our projections show that areas of endemism of five taxonomic groups (vascular plants, snails, spiders, butterflies, and beetles) in the Austrian Alps will, on average, experience a 77% habitat loss even under the weakest climate change scenario (+1.8 °C by 2100). The amount of habitat loss is positively related with the pooled endemic species richness (species from all five taxonomic groups) and with the richness of endemic vascular plants, snails, and beetles. Owing to limited postglacial migration, hotspots of high-altitude endemics are situated in rather low peripheral mountain chains of the Alps, which have not been glaciated during the Pleistocene. There, tree line expansion disproportionally reduces habitats of high-altitude species. Such legacies of climate history, which may aggravate extinction risks under future climate change have to be expected for many temperate mountain ranges.
Forest responses to climate change will depend on demographic impacts in the context of competition. Current models used to predict species responses, termed climate envelope models (CEMs), are controversial, because (i) calibration and prediction are based on correlations in space (CIS) between species abundance and climate, rather than responses to climate change over time (COT), and (ii) they omit competition. To determine the relative importance of COT, CIS, and competition for light, we applied a longitudinal analysis of 27 000 individual trees over 6–18 years subjected to experimental and natural variation in risk factors. Sensitivities and climate and resource tracking identify which species are vulnerable to these risk factors and in what ways. Results show that responses to COT differ from those predicted based on CIS. The most important impact is the effect of spring temperature on fecundity, rather than any input variable on growth or survival. Of secondary importance is growing season moisture. Species in the genera Pinus, Ulmus, Magnolia, and Fagus are particularly vulnerable to climate variation. However, the effect of competition on growth and mortality risk exceeds the effects of climate variation in space or time for most species. Because sensitivities to COT and competition are larger than CIS, current models miss the most important effects. By directly comparing sensitivity to climate in time and space, together with competition, the approach identifies which species are sensitive to climate change and why, including the heretofore overlooked impact on fecundity.
Studies on Fagus sylvatica show that growth in populations toward the southern limit of this species' distribution is limited strongly by drought. Warming temperatures in the Mediterranean region are expected to exacerbate drought where they are not accompanied by increases in precipitation. We studied levels of annual growth in mature F. sylvatica trees over the last half-century in the Montseny Mountains in Catalonia (northeast Spain). Our results show significantly lower growth of mature trees at the lower limit of this species' distribution when compared with trees at higher altitudes. Growth at the lower Fagus limit is characterized by a rapid recent decline starting in approximately 1975. By 2003, growth of mature trees had fallen by 49% when compared with predecline levels. This is not an age-related phenomenon, nor is it seen in comparable populations at higher altitudes. Analysis of climate-growth relationships suggests that the observed decline in growth is a result of warming temperatures and that, as precipitation in the region has not increased, precipitation is now insufficient to ameliorate the negative effects of increased temperatures on tree growth. As the climate-response of the studied forest is comparable with that of F. sylvatica forests in other southern European regions, it is possible that this growth decline is a more widespread phenomenon. Warming temperatures may lead to a rapid decline in the growth of range-edge populations and a consequent retreat of the species distribution in southern Europe. Assessment of long-term growth trends across the southern range edge of F. sylvatica therefore merits further attention.
Summary 1. Understanding the factors influencing tree growth is central to forest ecology because of the significance of growth to forest structure and biomass. One of the simplest, yet most controversial growth models, proposed by Enquist and colleagues, predicts that stem-diameter growth scales as the one-third power of stem diameter. Recent analyses of large-scale data sets have challenged the generality of this theory and highlighted the influence of resource competition on the scaling of growth with size. 2. Here we explore the factors regulating the diameter growth of 3334 trees of mountain beech ( Nothofagus solandri var. cliffortioides ) growing in natural single-species forests in New Zealand. Maximum-likelihood modelling was used to quantify the influences of tree size, altitude, the basal area of taller neighbours ( B L ) and the basal area of all neighbours ( B T ) on growth. Our interpretation of the models assumed that taller neighbours compete for light whereas all neighbours compete for nutrients. 3. The regression analyses indicate that competition for light has a strong influence on the growth of small trees, whereas competition for nutrients affects trees of all sizes. These findings are consistent with experimental manipulation studies showing that competition for light and nutrients inhibits the growth of small mountain beech trees, and fertilizer application studies showing that nitrogen limits the growth of large trees. 4. Tree growth declined with altitude. The regression analyses suggest that the intensity of light competition also declines with altitude, when trees with similar B T and B L values were compared along the gradient. These results are consistent with observations that trees become stunted and have more open canopies at high altitudes. 5. Our study is the first to build the effects of competition and environment into Enquist's model of tree growth. We show that competitive interactions alter the scaling of mean growth rate with size, whereas altitude does not influence the scaling of potential growth rate with size.
In this paper, we have reviewed how the hydraulic design of trees influences the movement of water from roots to leaves. The hydraulic architecture of trees can limit their water relations, gas exchange throughout the crown of trees, the distribution of trees over different habitats and, perhaps, even tbe maximum height that a particular species can achieve. Parameters of particular importance include:
(1) tbe vulnerabihty of stems to drought induced cavitation events because cavitation reduces the hydraulic conductance of stems,
(2) the leaf specific conductivity of stems because it determines the pressure gradients and most negative water potentials needed to sustain evaporation from leaves,
(3) the water storage capacity of tissues because this might determine the ability of trees to survive long drought periods.
All of these parameters are determined by the structure and function of anatomical components of trees. Some of the ecological and physiological trade-offs of specific structures are discussed.
The wide size range of conifer tracheids and angiosperm vessels has important consequences for function. In both conduit types, bigger is better for conducting efficiency. The gain in efficiency with size is maximized by the control of conduit shape, which balances end-wall and lumen resistances. Although vessels are an order of magnitude longer than tracheids of the same diameter, they are not necessarily more efficient because they lack the low end-wall resistance of tracheids with torus-margo pits. Instead, vessels gain conducting efficiency over tracheids by achieving wider maximum diameters. End-walls contributed 56-64% to total xylem resistance in both conduit types, indicating that length limits conducting efficiency. Tracheid dimensions may be more limited by unicellularity and the need to supply strength to homoxylous wood than by the need to protect against cavitation. In contrast, the greater size of the multicellular vessel is facilitated by fibers that strengthen heteroxylous wood. Vessel dimensions may be most limited by the need to restrict intervessel pitting and cavitation by air-seeding. Stressful habitats that promote narrow vessels should favor coexistence of conifers and angiosperms. The evolution of vessels in angiosperm wood may have required early angiosperms to survive a phase of mechanic and hydraulic instability.
A variable span smoother (No. LCS-TR-5). Stanford Univ CA lab for computational statistics
- J H Friedman
Friedman, J. H. (1984). A variable span smoother (No. LCS-TR-5). Stanford Univ CA lab for computational statistics.