Adaptation and plasticity in aboveground allometry variation of four pine species along environmental gradients

Plant species aboveground allometry can be viewed as a functional trait that reflects the evolutionary trade‐off between above‐ and belowground resources. In forest trees, allometry is related to productivity and resilience in different environments, and it is tightly connected with a compromise between efficiency‐safety and competitive ability. A better understanding on how this trait varies within and across species is critical to determine the potential of a species/population to perform along environmental gradients. We followed a hierarchical framework to assess tree height‐diameter allometry variation within and across four common European Pinus species. Tree height‐diameter allometry variation was a function of solely genetic components –approximated by either population effects or clinal geographic responses of the population's site of origin– and differential genetic plastic responses –approximated by the interaction between populations and two climatic variables of the growing sites (temperature and precipitation)–. Our results suggest that, at the species level, climate of the growing sites set the tree height‐diameter allometry of xeric and mesic species (Pinus halepensis, P. pinaster and P. nigra) apart from the boreal species (P. sylvestris), suggesting a weak signal of their phylogenies in the tree height‐diameter allometry variation. Moreover, accounting for interpopulation variability within species for the four pine species aided to: (1) detect genetic differences among populations in allometry variation, which in P. nigra and P. pinaster were linked to gene pools –genetic diversity measurements–; (2) reveal the presence of differential genetic variation in plastic responses along two climatic gradients in tree allometry variation. In P. sylvestris and P. nigra, genetic variation was the result of adaptive patterns to climate, while in P. pinaster and P. halepensis, this signal was either weaker or absent, respectively; and (3) detect local adaptation in the exponent of the tree height‐diameter allometry relationship in two of the four species (P. sylvestris and P. nigra), as it was a function of populations' latitude and altitude variables. Our findings suggest that the four species have been subjected to different historical and climatic constraints that might have driven their aboveground allometry and promoted different life strategies.     

Phylogenetic assemblage structure of North American trees is more strongly shaped by glacial–interglacial climate variability in gymnosperms than in angiosperms

How fast does biodiversity respond to climate change? The relationship of past and current climate with phylogenetic assemblage structure helps us to understand this question. Studies of angiosperm tree diversity in North America have already suggested effects of current water–energy balance and tropical niche conservatism. However, the role of glacial–interglacial climate variability remains to be determined, and little is known about any of these relationships for gymnosperms. Moreover, phylogenetic endemism, the concentration of unique lineages in restricted ranges, may also be related to glacial–interglacial climate variability and needs more attention. We used a refined phylogeny of both angiosperms and gymnosperms to map phylogenetic diversity, clustering and endemism of North American trees in 100‐km grid cells, and climate change velocity since Last Glacial Maximum together with postglacial accessibility to recolonization to quantify glacial–interglacial climate variability. We found: (1) Current climate is the dominant factor explaining the overall patterns, with more clustered angiosperm assemblages toward lower temperature, consistent with tropical niche conservatism. (2) Long‐term climate stability is associated with higher angiosperm endemism, while higher postglacial accessibility is linked to to more phylogenetic clustering and endemism in gymnosperms. (3) Factors linked to glacial–interglacial climate change have stronger effects on gymnosperms than on angiosperms. These results suggest that paleoclimate legacies supplement current climate in shaping phylogenetic patterns in North American trees, and especially so for gymnosperms.     

Phenology and fecundity in 11 sympatric pioneer species of Macaranga (Euphorbiaceae)in Borneo

Reproductive traits of tropical tree species vary predictably in relation to successional stage, but this variation may be due to the species' phylogenetic histories rather than selective pressures imposed by regeneration requirements. Reproductive phenology, tree size at the onset of reproduction, and fecundity of 11 sympatric, closely related Macaranga species were studied to investigate within-species variation in reproductive traits in relation to resource availability, and among-species variation in relation to other life-history traits (shade tolerance, seed size and maximum tree size, H(max)) and consequently the requirements for forest-gap colonization. Nine species reproduced in synchronous episodes, and two species reproduced continuously over 32 mo. Episodic reproduction was most intense in 1992 following a severe drought. For several species, reproductive trees had greater light availability, lower fecundity in lower light levels, and lower growth rates than nonreproductive trees, reflecting resource-limited reproduction. Among species, H(max) was negatively correlated with shade tolerance and seed size. Tree size at the onset of reproduction and fecundity was strongly linked to this axis of life-history variation, but phenological pattern was not. Absolute tree size at the onset of reproduction was positively correlated with H(max) and negatively correlated with shade tolerance. Relative size at reproductive onset was not correlated with shade tolerance or H(max). Fecundity ranged four orders of magnitude among species and was correlated positively with H(max) and negatively with seed size and shade tolerance. The interrelationships among these reproductive and other life-history traits are strongly correlated with the species' requirements for gap colonization.     

Evolutionary and ecological causes of species richness patterns in North American angiosperm trees

Climate and evolutionary factors (e.g. diversification, time‐for‐speciation, niche conservatism) are both thought to be major drivers of species richness in regional assemblages. However, few studies have simultaneously investigated the relative effects of climate and evolutionary factors on species richness across a broad geographical extent. Here, we assess their relative effects on species richness of angiosperm trees across North America. Species richness of angiosperm trees in 1175 regional assemblages were related to climate and phylogenetic structure using a structural equation modeling (SEM) approach. Climate was quantified based on the mean temperature of the coldest month and mean annual precipitation. Evolutionary factors (time‐for‐speciation vs diversification) were inferred from phylogeny‐based measures of mean root distance, phylogenetic species variability, and net relatedness index. We found that at the continental scale, species richness is correlated with temperature and precipitation with approximately similar strength. In the SEM with net relatedness index and phylogenetic species variability and with all the 1175 quadrats, the total direct effect size of phylogenetic structure on species richness is greater than the total direct effect size of climate on species richness by a factor of 3.7. The specific patterns of phylogenetic structure (i.e. greater phylogenetic distances in more species rich regions) are consistent with the idea that time and niche conservatism drive richness patterns in North American angiosperm trees. We conclude that angiosperm tree species richness in regional assemblages in North America is more strongly related to patterns of phylogenetic relatedness than to climatic variation. The results of the present study support the idea that climatic and evolutionary explanations for richness patterns are not in conflict, and that evolutionary processes explain both the relationship between climate and richness and substantial variation in richness that is independent of climate.     

Trait syndromes among North American trees are evolutionarily conserved and show adaptive value over broad geographic scales

Adaptive syndromes and their evolutionary constraints represent a powerful construct for understanding plant distributions. However, it is unclear how the species requirements to face multiple stressors promotes syndrome formation and to which abiotic stressors these syndromes show adaptive value over broad geographic scales. We combined local occurrence data from the U.S. Forest Inventory and Analysis (FIA) of 219 angiosperm and 85 gymnosperm species living across the conterminous US with phylogenies and trait data to identify tree syndromes, their evolutionary conservatism, and their adaptive value over broad scales. Factor analyses and evolutionary model selection revealed that trees possess functional trait syndromes that are strongly conserved. Major syndromes at the species level differed between angiosperms and gymnosperms. While the two main syndromes in angiosperms were related to cold and drought‐waterlogging tolerance, in gymnosperms a trade‐off between shade and drought tolerance was the main syndrome followed by a growth‐fire resistance syndrome. Additional RLQ and fourth‐corner approaches revealed that trait syndromes at the community level were broadly similar to those observed at the species level for angiosperms, although this was less clear for gymnosperms. This suggests that syndrome evolution has played an important role on angiosperm distributions, whereas additional ecological factors explain gymnosperm distributions. Importantly, syndromes show adaptive value, as they were geographically associated with several environmental variables showing structure from continental to local scales, being temperature the main abiotic stressor. Our results indicate that across the conterminous US tree species possess clear syndromes that are subjected to strong evolutionary constraints driving tree species and forest community distribution.     

Environmental correlates for tropical tree diversity and distribution patterns in Borneo

Aim Identify environmental correlates for tropical tree diversity and composition. Location Borneo, Southeast Asia. Methods A GIS‐environmental database with 5 arc minute (c. 10 × 10 km) resolution was combined with tree inventory data. Tree diversity, phylogenetic diversity (PD) and the two main compositional gradients were determined for 46 tree inventories. Akaike's information criterion and a data jackknifing procedure were used to select 50 explanatory models for diversity and composition gradients. The average of these models was used as our final diversity and compositional model. We applied Moran's I to detect spatial autocorrelation of residuals. Results Tree diversity, PD and the two main compositional gradients in Borneo were all significantly correlated with the environment. Tree diversity correlated negatively with elevation, soil depth, soil coarseness (texture) and organic carbon content, whereas it correlated positively with soil C:N ratio, soil pH, moisture storage capacity and annual rainfall. Tree PD was correlated positively with elevation and temperature seasonality and was largely determined by gymnosperms. However, angiosperm PD also correlated positive with elevation. Compositional patterns were strongly correlated with elevation but soil texture, cation‐exchange‐capacity, C:N ratio, C and N content and drainage were also important next to rainfall seasonality and El Niño Southern Oscillation drought impact. Main conclusions Although elevation is the most important correlate for diversity and compositional gradients in Borneo, significant additional variability is explained by soil characteristics (texture, carbon content, pH, depth, drainage and nutrient status) and climate (annual rainfall, rainfall seasonality and droughts). The identified environmental correlates for diversity and composition gradients correspond to those found in other tropical regions of the world. Differences between the regions are mainly formed by differences in the relative importance of the environmental variables in explaining diversity and compositional gradients.     

Phylogenetic properties of Tertiary relict flora in the east Asian continental islands: imprint of climatic niche conservatism and in situ diversification

Understanding biodiversity patterns on islands has long been a central aim in ecology and conservation biology. Island‐specific biogeographical processes play substantial roles in the formation of endemic biota. Here, we examined how climate niche conservatism and geohistorical factors are interactively associated with in situ diversification of Tertiary relict flora in the east Asian continental islands. We generated two novel datasets for species distribution and phylogeny that included all of the known vascular plant species in Japan (5575). Then we tested phylogenetic signal of climatic tolerance, in terms of absolute minimum temperature and water balance, and explored environmental predictors of phylogenetic structure (evolutionary derivedness and clustering) of species assemblages. Although phylogenetic signal of climatic tolerance was significant across the phylogeny of most species, the strength of climatic niche conservatism differed among ferns, gymnosperms, angiosperm trees, and angiosperm herbs. For angiosperm trees, cold temperatures acted as environmental filters that generated phylogenetic derivedness/clustering of species assemblages. For fern and angiosperm herb species, however, phylogenetic properties were not associated with climatic harshness. These contrasting patterns among groups reflected climate niche evolution in vascular plants with different growth forms and traits; for example, diversification of angiosperm trees (but not fern and herb) occurred in response to historical climatic cooling. More importantly, geographical constraints contributed to evolutionary radiation that resulted from isolation by distance from the continent or by elevation. Quaternary climate change was also associated with clade‐specific radiation in refugial habitats. The degree to which geographical, geological, and palaeoclimatic variables explain the phylogenetic structure underscores the importance of isolation‐ and habitat‐stability‐related geohistorical processes in driving in situ diversification despite climatic niche conservatism. We propose that the highly endemic flora of the east Asian islands resulted from the interplay of idiosyncratic regional factors, and ecological and evolutionary processes, such as climate niche assembly and adaptive/nonadaptive radiation.     

Predicting the height of fossil plant remains: an allometric approach to an old problem

Interspecific correlations between plant height and basal stem diameter (the allometry of height) and between stem length distal to where diameter is measured (the allometry of stem taper) were determined for a total of 265 species with self-supporting stems and for the shoots of five conifers and 15 angiosperm vine species. The allometric equations obtained for these data are proposed as a method to predict the heights of fossil plants for which basal stem diameters are either known or inferred, and to predict the missing lengths of fragmented stems based on the most proximal stem diameters observed.     

Crown allometries are less responsive than stem allometry to tree size and habitat variations in an Indian monsoon forest

While theoretical allometric models postulate universal scaling exponents, empirical relationships between tree dimensions show marked variability that reflects changes in the biomass allocation pattern. As growth of the various tree compartments may be controlled by different functions, it is hypothesized that they may respond differently to factors of variation, resulting in variable tree morphologies and potentially in trade-offs between allometric relationships. We explore the variability of tree stem and crown allometries using a dataset of 1,729 trees located in an undisturbed wet evergreen forest of the Western Ghats, India. We specifically test whether species adult stature, terrain slope, tree size and crown light exposure affect the relationships between stem diameter and stem height (stem allometry), and between stem diameter and crown width, crown area and crown volume (crown allometries). Results show that both stem and crown allometries are subject to variations in relation to both endogenous (tree size, species adult stature) and exogenous (terrain slope, crown light exposure) factors. Stem allometry appears to be more affected by these factors than are crown allometries, including the stem diameter–crown volume relationship, which proved to be particularly stable. Our results support the idea that height is a prevailing adjustment factor for a tree facing variable growth (notably light) conditions, while stem diameter–crown volume allometry responds more to internal metabolic constraints. We ultimately discuss the various sources of variability in the stem and crown allometries of tropical trees that likely play an important role in forest community dynamics.     

Evidence of variant intra- and interspecific scaling of tree crown structure and relevance for allometric theory

General scaling rules or constants for metabolic and structural plant allometry as assumed by the theory of Euclidian geometric scaling (2/3-scaling) or metabolic scaling (3/4-scaling) may meet human's innate propensity for simplicity and generality of pattern and processes in nature. However, numerous empirical works show that variability of crown structure rather than constancy is essential for a tree's success in coping with crowding. In order to link theory and empiricism, we analyzed the intra- and inter-specific scaling of crown structure for 52 tree species. The basis is data from 84 long-term plots of temperate monospecific forests under survey since 1870 and a set of 126 yield tables of angiosperm and gymnosperm forest tree species across the world. The study draws attention to (1) the intra-specific variation and correlation of the three scaling relationships: tree height versus trunk diameter, crown cross-sectional area versus trunk diameter, and tree volume versus trunk diameter, and their dependence on competition, (2) the inter-specific variation and correlation of the same scaling exponents ([Formula: see text] and [Formula: see text]) across 52 tree species, and (3) the relevance of the revealed variable scaling of crown structure for leaf organs and metabolic scaling. Our results arrive at suggesting a more extended metabolic theory of ecology which includes variability and covariation between allometric relationships as prerequisite for the individual plant's competitiveness.     

Crown development in a pioneer tree, Rhus trichocarpa, in relation to the structure and growth of individual branches

Based on an allometric reconstruction, the structure and biomass-allocation patterns of branches and current-year shoots were investigated in branches of various heights in the pioneer tree Rhus trichocarpa, to evaluate how crown development is achieved and limited in association with height. Path analysis was conducted to explore the effects of light availability, basal height and size of individual branches on branch structure and growth. Branch angle was affected by basal height, whereas branch mass was influenced primarily by light availability. This result suggests that branch structure is strongly constrained by basal height, and that trees mediate such constraints under different light environments. Previous-year leaf area and light availability showed positive effects on current-year stem mass. In contrast, branch basal height and mass negatively affected current-year stem mass. Moreover, the length of stems of a given diameter decreased with increasing branch height. Therefore the cost of biomass investment for a unit growth in length is greater for branches of larger size and at upper positions. Vertical growth rate in length decreased with increasing height. Height-dependent changes in stem allometry and angle influenced the reduction in vertical growth rate to a similar degree.     

Latitudinal Variation in Leaf and Tree Traits of the Mangrove Avicennia germinans (Avicenniaceae) in the Central Region of the Gulf of Mexico

The variation in leaf mass per area, leaf nutrients (% carbon, nitrogen and phosphorus), and the allometric relation between tree height and diameter of the black mangrove, Avicennia germinans, were explored in nine mangrove forests in similar environments along a 5° latitudinal gradient in the central region of the Gulf of Mexico, as indicated by a southward increase in temperature and precipitation. There was no correlation between leaf nitrogen or phosphorus content and latitude. Leaf mass per area and leaf carbon content were positively correlated with latitude and negatively correlated with temperature and annual rainfall, whereas asymptotic tree height and maximum diameter showed the opposite trend. Such patterns suggest a trade-off between leaf traits and tree size which may be constrained by the same environmental factors along a dry-cold to humid-warm latitudinal gradient.     

Climate‐ and successional‐related changes in functional composition of European forests are strongly driven by tree mortality

Intense droughts combined with increased temperatures are one of the major threats to forest persistence in the 21st century. Despite the direct impact of climate change on forest growth and shifts in species abundance, the effect of altered demography on changes in the composition of functional traits is not well known. We sought to (1) quantify the recent changes in functional composition of European forests; (2) identify the relative importance of climate change, mean climate and forest development for changes in functional composition; and (3) analyse the roles of tree mortality and growth underlying any functional changes in different forest types. We quantified changes in functional composition from the 1980s to the 2000s across Europe by two dimensions of functional trait variation: the first dimension was mainly related to changes in leaf mass per area and wood density (partially related to the trait differences between angiosperms and gymnosperms), and the second dimension was related to changes in maximum tree height. Our results indicate that climate change and mean climatic effects strongly interacted with forest development and it was not possible to completely disentangle their effects. Where recent climate change was not too extreme, the patterns of functional change generally followed the expected patterns under secondary succession (e.g. towards late‐successional short‐statured hardwoods in Mediterranean forests and taller gymnosperms in boreal forests) and latitudinal gradients (e.g. larger proportion of gymnosperm‐like strategies at low water availability in forests formerly dominated by broad‐leaved deciduous species). Recent climate change generally favoured the dominance of angiosperm‐like related traits under increased temperature and intense droughts. Our results show functional composition changes over relatively short time scales in European forests. These changes are largely determined by tree mortality, which should be further investigated and modelled to adequately predict the impacts of climate change on forest function.     

Treeline dynamics in relation to climatic variability in the central Tianshan Mountains, northwestern China

Aim Climate variability may be an important mediating agent of ecosystem dynamics in cold, arid regions such as the central Tianshan Mountains, north‐western China. Tree‐ring chronologies and the age structure of a Schrenk spruce (Picea schrenkiana) forest were developed to examine treeline dynamics in recent decades in relation to climatic variability. Of particular interest was whether tree‐ring growth and population recruitment patterns responded similarly to climate warming. Location The study was conducted in eight stands that ranged from 2500 m to 2750 m a.s.l. near the treeline in the Tianchi Nature Reserve (43°45′?43°59′ N, 88°00′?88°20′ E) in the central Xinjiang Uygur Autonomous Region, northwestern China. Methods Tree‐ring cores were collected and used to develop tree‐ring chronologies. The age of sampled trees was determined from basal cores sampled as close as possible to the ground. Population age structure and recruitment information were obtained using an age–d.b.h. (diameter at breast height) regression from the sampled cores and the d.b.h. measured on all trees in the plots. Ring‐width chronologies and tree age structure were both used to investigate the relationship between treeline dynamics and climate change. Results Comparisons with the climatic records showed that both the radial growth of trees and tree recruitment were influenced positively by temperature and precipitation in the cold high treeline zone, but the patterns of their responses differed. The annual variation in tree rings could be explained largely by the average monthly minimum temperatures during February and August of the current year and by the monthly precipitation of the previous August and January, which had a significant and positive effect on tree radial growth. P. schrenkiana recruitment was influenced mainly by consecutive years of high minimum summer temperatures and high precipitation during spring. Over the last several decades, the treeline did not show an obvious upward shift and new recruitment was rare. Some trees had established at the treeline at least 200 years ago. Recruitment increased until the early 20th century (1910s) but then decreased with poor recruitment over the past several decades (1950–2000). Main conclusions There were strong associations between climatic change and ring‐width patterns, and with recruitments in Schrenk spruce. Average minimum temperatures in February and August, and total precipitation in the previous August and January, had a positive effect on tree‐ring width, and several consecutive years of high minimum summer temperature and spring precipitation was a main factor favouring the establishment of P. schrenkiana following germination within the treeline ecotone. Both dendroclimatology and recruitment analysis were useful and compatible to understand and reconstruct treeline dynamics in the central Tianshan Mountains.     

Dual impacts of climate change: forest migration and turnover through life history

Tree species are predicted to track future climate by shifting their geographic distributions, but climate‐mediated migrations are not apparent in a recent continental‐scale analysis. To better understand the mechanisms of a possible migration lag, we analyzed relative recruitment patterns by comparing juvenile and adult tree abundances in climate space. One would expect relative recruitment to be higher in cold and dry climates as a result of tree migration with juveniles located further poleward than adults. Alternatively, relative recruitment could be higher in warm and wet climates as a result of higher tree population turnover with increased temperature and precipitation. Using the USDA Forest Service's Forest Inventory and Analysis data at regional scales, we jointly modeled juvenile and adult abundance distributions for 65 tree species in climate space of the eastern United States. We directly compared the optimal climate conditions for juveniles and adults, identified the climates where each species has high relative recruitment, and synthesized relative recruitment patterns across species. Results suggest that for 77% and 83% of the tree species, juveniles have higher optimal temperature and optimal precipitation, respectively, than adults. Across species, the relative recruitment pattern is dominated by relatively more abundant juveniles than adults in warm and wet climates. These different abundance‐climate responses through life history are consistent with faster population turnover and inconsistent with the geographic trend of large‐scale tree migration. Taken together, this juvenile–adult analysis suggests that tree species might respond to climate change by having faster turnover as dynamics accelerate with longer growing seasons and higher temperatures, before there is evidence of poleward migration at biogeographic scales.     

Adaptive and plastic responses of Quercus petraea populations to climate across Europe

How temperate forests will respond to climate change is uncertain; projections range from severe decline to increased growth. We conducted field tests of sessile oak (Quercus petraea), a widespread keystone European forest tree species, including more than 150 000 trees sourced from 116 geographically diverse populations. The tests were planted on 23 field sites in six European countries, in order to expose them to a wide range of climates, including sites reflecting future warmer and drier climates. By assessing tree height and survival, our objectives were twofold: (i) to identify the source of differential population responses to climate (genetic differentiation due to past divergent climatic selection vs. plastic responses to ongoing climate change) and (ii) to explore which climatic variables (temperature or precipitation) trigger the population responses. Tree growth and survival were modeled for contemporary climate and then projected using data from four regional climate models for years 2071–2100, using two greenhouse gas concentration trajectory scenarios each. Overall, results indicated a moderate response of tree height and survival to climate variation, with changes in dryness (either annual or during the growing season) explaining the major part of the response. While, on average, populations exhibited local adaptation, there was significant clinal population differentiation for height growth with winter temperature at the site of origin. The most moderate climate model (HIRHAM5‐EC; rcp4.5) predicted minor decreases in height and survival, while the most extreme model (CCLM4‐GEM2‐ES; rcp8.5) predicted large decreases in survival and growth for southern and southeastern edge populations (Hungary and Turkey). Other nonmarginal populations with continental climates were predicted to be severely and negatively affected (Bercé, France), while populations at the contemporary northern limit (colder and humid maritime regions; Denmark and Norway) will probably not show large changes in growth and survival in response to climate change.     

Floristic and functional affiliations of woody plants with climate in western Amazonia

Aim To test whether a direct relationship exists between the relative abundance of woody plant genera and precipitation regime along the north–south climate gradient of the western Amazon. Location Lowland rain forests in the western Amazon. Methods Floristic data on 91 woody plant genera, from 39 0.1‐ha plots across the western Amazon, and precipitation data from a 0.5° global data set were used to test for correlations between plant relative abundance (defined as percentage number of stems ≥ 2.5 cm diameter at breast height for each woody plant genus per plot) and derived dry‐season variables. Moisture preference was then assessed in terms of pioneer and shade‐tolerant life‐history strategy. Results There were significant associations between the distribution of plant relative abundances and seasonal precipitation variables in 34% of genera analysed. Significant differences were identified in size‐class distribution between dry affiliates and generalists. Dry affiliates were not dominant in any size class in any plot type, whereas climate generalists dominated most of the size classes in the dry plots and the mid‐range size classes in the wet plots. Dry‐affiliate genera were a minority, even in dry forests. Wet‐affiliate genera were correlated with shade tolerance, whereas genera with no rainfall affiliation were often pioneers. Main conclusions The results suggest that moisture variable seasonality influences community composition in a manner that can be related to the life‐history trade‐off between shade tolerance and pioneer ranking. One possible reason for higher diversity in wetter forests is that high rainfall amplifies the niche space available to shade‐tolerant plants. Determining which plant groups are constrained by which environmental variables can contribute to our understanding of how forest composition may be changing now, and how it may change under future climate: if shade‐tolerant trees are also drought‐intolerant, community structure in wet forests may be more vulnerable to future increases in moisture stress.     

Linking litter calcium, earthworms and soil properties: a common garden test with 14 tree species

Tree species can influence biogeochemistry through variation in the quantity and chemistry of their litter, and associated impacts on the soil heterotrophic community. However, the role that different plant traits play in these processes is not well understood, nor is it clear whether species effects on soils largely reflect a gymnosperm vs. angiosperm contrast. Using a replicated, long‐term monoculture plot experiment, we examined variation in soils among 14 gymnosperm and angiosperm tree species 30 years after plot establishment, and assessed the role of litter chemistry vis‐à‐vis such variation. Differences in litter calcium concentrations among tree species resulted in profound changes in soil acidity and fertility that were similar within and among tree groups. Tree species rich in calcium were associated with increased native earthworm abundance and diversity, as well as increased soil pH, exchangeable calcium, per cent base saturation and forest floor turnover rate.     

Architecture of Iberian canopy tree species in relation to wood density,shade tolerance and climate

Tree architecture has important consequences for tree performance as it determines resource capture, mechanical stability and dominance over competitors. We analyzed architectural relationships between stem and crown dimensions for 13 dominant Iberian canopy tree species belonging to the Pinaceae (six Pinus species) and Fagaceae (six Quercus species and Fagus sylvatica) and related these architectural traits to wood density, shade tolerance and climatic factors. Fagaceae had, compared with Pinaceae, denser wood, saplings with wider crowns and adults with larger maximal crown size but smaller maximal height. In combination, these traits enhance light acquisition and persistence in shaded environments; thus, contributing to their shade tolerance. Pinaceae species, in contrast, had low-density wood, allocate more resources to the formation of the central trunk rather than to branches and attained taller maximal heights, allowing them to grow rapidly in height and compete for light following disturbances; thus, contributing to their high light requirements. Wood density had a strong relationship with tree architecture, with dense-wooded species having smaller maximum height and wider crowns, probably because of cheaper expansion costs for producing biomechanically stable branches. Species from arid environments had shorter stems and shallower crowns for a given stem diameter, probably to reduce hydraulic path length and assure water transport. Wood density is an important correlate of variation in tree architecture between species and the two dominant families, with potentially large implications for their resource foraging strategies and successional dynamics.     

Phylogenetic structure of angiosperm trees in local forest communities along latitudinal and elevational gradients in eastern North America

Latitudinal and elevational gradients both represent thermal gradients. Assessing the consistency of the relationships between phylogenetic structure and climate between latitudinal and elevational gradients can provide insight into the mechanisms driving assembly of species from regional pools into local assemblages. The aim of this study is to compare patterns of phylogenetic structure measures for angiosperm tree species between latitudinal and elevational gradients, using a dataset of angiosperm tree species in 14 092 forest plots in eastern North America. We assessed whether these two gradients produce similar relationships between climate and phylogenetic structure, hypothesizing that they should differ in magnitude but not direction. We used correlation and regression analyses to assess the relation of measures of phylogenetic structure to elevation, latitude and climatic variables, which included minimum temperature, temperature seasonality, annual precipitation and precipitation seasonality. We found that 1) phylogenetic relatedness of angiosperm trees increases with decreasing temperature along both latitudinal and elevational gradients but the relationship between phylogenetic relatedness and temperature is steeper for elevational gradients than for latitudinal gradients; 2) the tip-weighted metric of phylogenetic relatedness (nearest taxon index) is more strongly correlated with climatic variables than the basal-weighted metric of phylogenetic relatedness (net relatedness index); 3) winter cold temperature exerts a stronger effect on community assembly of angiosperm trees than does temperature seasonality. These results suggest that winter cold temperature, rather than temperature seasonality, drives phylogenetic structure of plants in local forest communities, and that species distributions along elevational gradients are more in equilibrium with temperature, compared with those along latitudinal gradients.