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Fundamental axes of variation in plant traits result from trade-offs between costs and benefits of resource-use strategies at the leaf scale. However, it is unclear whether similar trade-offs propagate to the ecosystem level. Here, we test whether trait correlation patterns predicted by three well-known leaf- and plant-level coordination theories - the leaf economics spectrum, the global spectrum of plant form and function, and the least-cost hypothesis - are also observed between community mean traits and ecosystem processes. We combined ecosystem functional properties from FLUXNET sites, vegetation properties, and community mean plant traits into three corresponding principal component analyses. We find that the leaf economics spectrum (90 sites), the global spectrum of plant form and function (89 sites), and the least-cost hypothesis (82 sites) all propagate at the ecosystem level. However, we also find evidence of additional scale-emergent properties. Evaluating the coordination of ecosystem functional properties may aid the development of more realistic global dynamic vegetation models with critical empirical data, reducing the uncertainty of climate change projections., (© 2023. The Author(s).)

As Earth's climate has varied strongly through geological time, studying the impacts of past climate change on biodiversity helps to understand the risks from future climate change. However, it remains unclear how paleoclimate shapes spatial variation in biodiversity. Here, we assessed the influence of Quaternary climate change on spatial dissimilarity in taxonomic, phylogenetic, and functional composition among neighboring 200-kilometer cells (beta-diversity) for angiosperm trees worldwide. We found that larger glacial-interglacial temperature change was strongly associated with lower spatial turnover (species replacements) and higher nestedness (richness changes) components of beta-diversity across all three biodiversity facets. Moreover, phylogenetic and functional turnover was lower and nestedness higher than random expectations based on taxonomic beta-diversity in regions that experienced large temperature change, reflecting phylogenetically and functionally selective processes in species replacement, extinction, and colonization during glacial-interglacial oscillations. Our results suggest that future human-driven climate change could cause local homogenization and reduction in taxonomic, phylogenetic, and functional diversity of angiosperm trees worldwide.

Safeguarding Earth's tree diversity is a conservation priority due to the importance of trees for biodiversity and ecosystem functions and services such as carbon sequestration. Here, we improve the foundation for effective conservation of global tree diversity by analyzing a recently developed database of tree species covering 46,752 species. We quantify range protection and anthropogenic pressures for each species and develop conservation priorities across taxonomic, phylogenetic, and functional diversity dimensions. We also assess the effectiveness of several influential proposed conservation prioritization frameworks to protect the top 17% and top 50% of tree priority areas. We find that an average of 50.2% of a tree species' range occurs in 110-km grid cells without any protected areas (PAs), with 6,377 small-range tree species fully unprotected, and that 83% of tree species experience nonnegligible human pressure across their range on average. Protecting high-priority areas for the top 17% and 50% priority thresholds would increase the average protected proportion of each tree species' range to 65.5% and 82.6%, respectively, leaving many fewer species (2,151 and 2,010) completely unprotected. The priority areas identified for trees match well to the Global 200 Ecoregions framework, revealing that priority areas for trees would in large part also optimize protection for terrestrial biodiversity overall. Based on range estimates for >46,000 tree species, our findings show that a large proportion of tree species receive limited protection by current PAs and are under substantial human pressure. Improved protection of biodiversity overall would also strongly benefit global tree diversity.

Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a wealth of studies exploring trait relationships across the entire plant kingdom, the dominant traits underpinning these unique aspects of tree form and function remain unclear. Here, by considering 18 functional traits, encompassing leaf, seed, bark, wood, crown, and root characteristics, we quantify the multidimensional relationships in tree trait expression. We find that nearly half of trait variation is captured by two axes: one reflecting leaf economics, the other reflecting tree size and competition for light. Yet these orthogonal axes reveal strong environmental convergence, exhibiting correlated responses to temperature, moisture, and elevation. By subsequently exploring multidimensional trait relationships, we show that the full dimensionality of trait space is captured by eight distinct clusters, each reflecting a unique aspect of tree form and function. Collectively, this work identifies a core set of traits needed to quantify global patterns in functional biodiversity, and it contributes to our fundamental understanding of the functioning of forests worldwide., (© 2022. The Author(s).)

Background: Several smartphone applications have been developed for the purpose of low-cost and convenient assessments of vegetation canopy structure and understorey illumination. Like standard hemispherical photography, most of these applications require user decisions about image processing, posing challenges for repeatability of measurements. Here I report a test of CanopyCapture, an application that instantaneously estimates percentage canopy gap fraction without any input from the user, and has the added advantage of an intuitive levelling mechanism., Methods: Gap fraction estimates by CanopyCapture were compared with gap fraction values computed by the LAI-2200C Canopy Analyzer, in two contrasting evergreen temperate forests in New Zealand: an even-aged southern beech ( Nothofagus ) stand and an old-growth podocarp/broadleaf forest. These comparisons were repeated using a wide-angle adapter to enhance the smartphone camera's field of view from 45 to 65°. I also asked if CanopyCapture results depended on sky condition (sunny vs . overcast) and on the type of smartphone used., Results: CanopyCapture output was significantly correlated with gap fraction computed by the LAI-2200C (R 2 = 0.39), and use of the wide-angle adapter lifted this value to 0.56. However, CanopyCapture output was not significantly correlated with LAI-2200C output in the even-aged Nothofagus stand, where there was less spatial variation in canopy structure. Despite being much less sensitive to variation in gap fraction than the LAI-2200C, CanopyCapture was nevertheless able to detect differences in average gap fraction between the two forests studied. CanopyCapture results beneath intact canopies were not significantly affected by sky condition, but reflection of direct light off tree trunks in sunny weather caused slight overestimation of gap fraction beneath broken canopies and gaps. Uneven or patchy cloud cover can also cause erroneous readings beneath large canopy openings. Three different models of smartphone gave different results., Conclusions: CanopyCapture offers a rapid and repeatable proxy for comparisons of average canopy gap fraction in multiple stands/forests, provided large sample sizes are used. Measurement under even overcast skies is recommended, and studies involving multiple operators will need to standardize smartphones to ensure comparability of results. Although wide-angle adapters can improve performance, CanopyCapture's low sensitivity prevents high-resolution comparisons of the light environments of individual understorey plants within a stand., Competing Interests: The author declares he has no competing interests., (© 2022 Lusk.)

Some plant traits may be legacies of coevolution with extinct megafauna. One example is the convergent evolution of 'divaricate' cage architectures in many New Zealand lineages, interpreted as a response to recently extinct flightless avian browsers whose ancestors arrived during the Paleogene period. Although experiments have confirmed that divaricate habit deters extant browsers, its abundance on frosty, droughty sites appears consistent with an earlier interpretation as a response to cold, dry Plio-Pleistocene climates. We used 45 protein-coding sequences from plastid genomes to reconstruct the evolutionary history of the divaricate habit in extant New Zealand lineages. Our dated phylogeny of 215 species included 91% of New Zealand eudicot divaricate species. We show that 86% of extant divaricate plants diverged from non-divaricate sisters within the last 5 Ma, implicating Plio-Pleistocene climates in the proliferation of cage architectures in New Zealand. Our results, combined with other recent findings, are consistent with the synthetic hypothesis that the browser-deterrent effect of cage architectures was strongly selected only when Plio-Pleistocene climatic constraints prevented woody plants from growing quickly out of reach of browsers. This is consistent with the abundance of cage architectures in other regions where plant growth is restricted by aridity or short frost-free periods., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

We introduce the AusTraits database - a compilation of values of plant traits for taxa in the Australian flora (hereafter AusTraits). AusTraits synthesises data on 448 traits across 28,640 taxa from field campaigns, published literature, taxonomic monographs, and individual taxon descriptions. Traits vary in scope from physiological measures of performance (e.g. photosynthetic gas exchange, water-use efficiency) to morphological attributes (e.g. leaf area, seed mass, plant height) which link to aspects of ecological variation. AusTraits contains curated and harmonised individual- and species-level measurements coupled to, where available, contextual information on site properties and experimental conditions. This article provides information on version 3.0.2 of AusTraits which contains data for 997,808 trait-by-taxon combinations. We envision AusTraits as an ongoing collaborative initiative for easily archiving and sharing trait data, which also provides a template for other national or regional initiatives globally to fill persistent gaps in trait knowledge., (© 2021. The Author(s).)

The abundance of the divaricate growth form in New Zealand has been interpreted as either (a) the response of an isolated flora to cool, dry, Plio-Pleistocene climates; or (b) a defense against large browsing birds (moa) that were hunted to extinction shortly after human arrival during the last millennium. We used patterns of divaricate plant abundance across present-day landscapes to test a novel synthetic hypothesis: that the divaricate form is of most value to plants on fertile soils that attract herbivores, on sites where climatic constraints prevent plants from quickly growing out of the browse zone. This hypothesis predicts that divaricate species should be most abundant on terraces (landforms that are both fertile and frost-prone) in regions that are cold and dry, and should be scarce across all topographic positions in the warmest (largely frost-free) regions. To address our hypothesis, we first tested the influence of topography on frost regimes and nutrient levels by measuring temperatures and soil total C, N, and P at four standard topographic positions at five localities differing widely in macroclimate. We then extracted a dataset of 236 surveys comprising 9,877 relevé plots from the New Zealand National Vegetation Survey databank. We calculated the proportion of arborescent species with a divaricate growth form and the proportion of total arborescentcover contributed by divaricates on each plot; we then fitted linear mixed-effect models predicting these response variables as functions of topographic position and climate. The number of frosts recorded averaged <1 yr -1 at the warmest of the five sites studied, to >60 yr -1 on all topographic positions at the coldest site. Terraces were subject to more frequent and harder frosts than any other topographic position. Topography had no significant influence on total N or C:N, but total P was higher on terraces and in gullies than on faces or ridges. Frost-free period was the dominant influence on both species representation and cover of divaricate plants throughout the country. The effect of topography was also significant, but weaker. The effect of frost-free period was stronger on sites with water deficits than on sites where precipitation exceeded evapotranspiration. Divaricates made their largest contributions on terraces in cold, dry regions; as predicted, they were scarce on all topographic positions on sites with frost-free periods >300 days. Our hypothesis was generally supported, although the effect of topography on divaricate abundance was not as strong as some previous studies led us to expect. Divaricates made their largest contributions to arborescent species richness and cover on sites where climatic restrictions on growth coincide with relatively high nutrient availability. The contemporary distribution of the divaricate form across New Zealand landscapes thus appears to be reasonably well explained by the hypothesized interaction of climate and fertility-mediated browsing, although experiments may provide more conclusive tests of this hypothesis., (Copyright © 2020 Lusk, Wiser and Laughlin.)

Leaf size varies conspicuously along environmental gradients. Small leaves help plants cope with drought and frost, because of the effect of leaf size on boundary layer conductance; it is less clear what advantage large leaves confer in benign environments. We asked if large leaves give species of warm climates an advantage in seedling light interception efficiency over small-leaved species from colder environments. We measured seedling leaf, architectural and biomass distribution traits of 18 New Zealand temperate rainforest evergreens; we then used a 3-D digitiser and the Yplant program to model leaf area display and light interception. Species associated with mild climates on average had larger leaves and larger specific leaf areas (SLA) than those from cold climates, and displayed larger effective foliage areas per unit of aboveground biomass, indicating higher light interception efficiency at whole-plant level. This reflected differences in total foliage area, rather than in self-shading. Our findings advance the understanding of leaf size by showing that large leaves enable seedlings of species with highly conductive (but frost-sensitive) xylem to deploy large foliage areas without increasing self-shading. Leaf size variation along temperature gradients in humid forests may therefore reflect a trade-off between seedling light interception efficiency and susceptibility to frost., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

The species-rich arborescent assemblages of humid tropical forests encompass much of the known range of the leaf economics spectrum, often including >20-fold variation in leaf lifespan. This suite of traits underpins a life-history continuum from fast-growing pioneers to slow-growing shade-tolerant species. Less is known about the range of leaf traits in humid temperate forests, and there are conflicting reports about relationships of these traits with the light requirements of temperate evergreen angiosperms. Here I quantify the range of leaf functional traits in a New Zealand temperate evergreen forest, and relationships of these traits with light requirements of juvenile trees and shrubs. Foliage turnover of saplings of 19 evergreen angiosperms growing beneath gaps (12-29% canopy openness) and in understories (1.2-2.9%) was measured over 12 months. Dry mass per area (LMA), dry matter content, thickness, density and nitrogen content (N) of leaves were also measured. Species minimum light requirements were indexed as the 10th percentile of the distribution of saplings in relation to canopy openness. Interspecific variation of leaf lifespan was ∼6-fold in gaps (0.6 to 3.8 yrs), and ∼11-fold in the understorey (0.7 to 7.7 yrs). Six small tree and shrub species are effectively leaf-exchangers, with leaf lifespans of c.1 year in gaps-albeit usually longer in the shade. Interspecific variation in other leaf traits was 2.5 to 4-fold. Lifespans and LMA of both sun and shade leaves were negatively correlated with species light requirements i.e., positively correlated with shade tolerance. However, light environment (gap vs shade) explained about the same amount of variation in LMA as species' identity did. Species light requirements were not significantly correlated with leaf N, dry matter content, density or thickness-except for a marginally significant correlation with dry matter content of shade leaves. Species light requirements were thus less consistently related to leaf structural traits than appears to be the case in humid tropical forests. Whereas the wide interspecific variation in leaf economic traits of tropical rainforest species outweighs plastic response to light availability, temperate evergreen woody angiosperms appear to occupy a narrower range of the leaf economic spectrum. Standardization of the light environments in which LMA is measured is vital in comparative studies of humid temperate forest evergreens, because of countergradient responses of this trait to light, and because of the relative magnitudes of plastic and interspecific variation in LMA in these forests., Competing Interests: The author declares there are no competing interests.

Explanations of leaf size variation commonly focus on water availability, yet leaf size also varies with latitude and elevation in environments where water is not strongly limiting. We provide the first conclusive test of a prediction of leaf energy balance theory that may explain this pattern: large leaves are more vulnerable to night-time chilling, because their thick boundary layers impede convective exchange with the surrounding air. Seedlings of 15 New Zealand evergreens spanning 12-fold variation in leaf width were exposed to clear night skies, and leaf temperatures were measured with thermocouples. We then used a global dataset to assess several climate variables as predictors of leaf size in forest assemblages. Leaf minus air temperature was strongly correlated with leaf width, ranging from -0.9 to -3.2°C in the smallest- and largest-leaved species, respectively. Mean annual temperature and frost-free period were good predictors of evergreen angiosperm leaf size in forest assemblages, but no climate variable predicted deciduous leaf size. Although winter deciduousness makes large leaves possible in strongly seasonal climates, large-leaved evergreens are largely confined to frost-free climates because of their susceptibility to radiative cooling. Evergreen leaf size data can therefore be used to enhance vegetation models, and to infer palaeotemperatures from fossil leaf assemblages., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

Recent work suggests that plant size affects light requirements and carbon balance of juvenile trees, and such shifts may be greater in light-demanding species than in their more shade-tolerant associates. To explore the physiological basis of such shifts, we measured juvenile light interception, carbon gain and growth of four subtropical Australian rainforest trees differing in shade tolerance, comparing individuals ranging from 13 to 238 cm in height, across a wide range of understory environments. We hypothesized that even in a standardized light environment, increasing sapling size would lead to declines in net daily carbon gain of foliage and relative growth rates (RGR) of all species, with declines more pronounced in light-demanding species. Crown architecture of individuals was recorded using a 3-dimensional digitizer, and the YPLANT program was used to estimate the self-shaded fraction of each crown and model net carbon gain. Increased sapling size caused a significant increase in self-shading, and significant declines in net daily carbon gain and RGR of light-demanding species, while such ontogenetic variations were minimal or absent in shade-tolerant species. Additionally, differences in the slope of the relationship between light and RGR led to crossovers in RGR among shade-tolerant and light-demanding species at low light. Our results show that the magnitude of ontogenetic variation in net daily carbon gain and RGR can be substantial and may depend on successional status, making it unsafe to assume that young seedling performance can be used to predict or model responses of larger juvenile trees.

The worldwide plant economic spectrum hypothesis predicts that leaf, stem, and root traits are correlated across vascular plant species because carbon gain depends on leaves being adequately supplied with water and nutrients, and because construction of each organ involves a trade-off between performance and persistence. Despite its logical and intuitive appeal, this hypothesis has received mixed empirical support. If traits within species diverge in their responses to an environmental gradient, then interspecific trait correlations could be weakened when measured in natural ecosystems. To test this prediction, we measured relative growth rates (RGR) and seven functional traits that have been shown to be related to fluxes of water, nutrients, and carbon across 56 functionally diverse tree species on (1) juveniles in a controlled environment, (2) juveniles in forest understories, and (3) mature trees in forests. Leaf, stem, and fine root traits of juveniles grown in a controlled environment were closely correlated with each other, and with RGR. Remarkably, the seven leaf, stem, and fine root tissue traits spanned a single dimension of variation when measured in the controlled environment. Forest-grown juveniles expressed lower leaf mass per area, but higher wood and fine root tissue density, than greenhouse-grown juveniles. Traits and growth rates were decoupled in forest-grown juveniles and mature trees. Our results indicate that constraints exist on the covariation, not just the variation, among vegetative plant organs; however, divergent responses of traits within species to environmental gradients can mask interspecific trait correlations in natural environments. Correlations among organs and relationships between traits and RGR were strong when plants were compared in a standardized environment. Our results may reconcile the discrepancies seen among studies, by showing that if traits and growth rates of species are compared across varied environments, then the interorgan trait correlations observed in controlled conditions can weaken or disappear.

Forest ecologists researching the functional basis of tree regeneration patterns and species coexistence often attempt to correlate traits with light-gradient partitioning. However, an exclusive focus on light can overlook other important drivers of forest dynamics. We measured light, temperatures, humidity and sapling densities in each of four phases of a forest dynamic mosaic in New Zealand: shaded understoreys, tree-fall gaps, treefern groves and clearings. We then measured leaf, wood and seed traits, as potential predictors of species' regeneration patterns. Saplings of 18 out of 21 species were significantly associated with one or other of the four phases, and associations were best predicted by a two-trait model (leaf size, wood density) explaining 51% of observed variation. Species associated with treefall gaps had traits favouring light pre-emption (large leaves, low-density wood), whereas those establishing in clearings mostly had small leaves and dense wood, traits probably conferring resistance to the frosts and summer water deficits that saplings were exposed to there. The dynamics of some forests cannot be explained adequately by light-gradient partitioning through a growth vs shade tolerance tradeoff, underpinned by the leaf economics spectrum. Consideration of multiple environmental filters and multiple traits will enhance understanding of regeneration patterns and species coexistence., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

Although shade tolerance is often assumed to be a fixed trait, recent work suggests ontogenetic changes in the light requirements of tree species. We determined the influence of gas exchange, biomass distribution, and self-shading on ontogenetic variation in the instantaneous aboveground carbon balance of Acer saccharum. We quantified the aboveground biomass distributions of 18 juveniles varying in height and growing in low light in a temperate forest understory in Minnesota, USA. Gas exchange rates of leaf and stem tissues were measured, and the crown architecture of each individual was quantified. The YPLANT program was used to estimate the self-shaded fraction of each crown and to model net leaf-level carbon gain. Leaf respiration and photosynthesis per gram of leaf tissue increased with plant size. In contrast, stem respiration rates per gram of stem tissue declined, reflecting a shift in the distribution of stem diameter sizes from smaller (with higher respiration) to larger diameter classes. However, these trends were outweighed by ontogenetic increases in self-shading (which reduces the net photosynthesis realized) and stem mass fraction (which increases the proportion of purely respiratory tissue) in terms of influence on net carbon exchange. As a result, net carbon gain per gram of aboveground plant tissue declined with increasing plant size, and the instantaneous aboveground light compensation point increased. When estimates of root respiration were included to model whole-plant carbon gain and light compensation points, relationships with plant size were even more pronounced. Our findings show how an interplay of gas exchange, self-shading, and biomass distribution shapes ontogenetic changes in shade tolerance.

Leaf dark respiration (Rdark ) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of Rdark and associated leaf traits. Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in Rdark . Area-based Rdark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8-28°C). By contrast, Rdark at a standard T (25°C, Rdark (25) ) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher Rdark (25) at a given photosynthetic capacity (Vcmax (25) ) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark (25) values at any given Vcmax (25) or [N] were higher in herbs than in woody plants. The results highlight variation in Rdark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of Rdark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs)., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Unlabelled: •, Premise of Study: Have Gondwanan rainforest floral associations survived? Where do they occur today? Have they survived continuously in particular locations? How significant is their living floristic signal? We revisit these classic questions in light of significant recent increases in relevant paleobotanical data.•, Methods: We traced the extinction and persistence of lineages and associations through the past across four now separated regions-Australia, New Zealand, Patagonia, and Antarctica-using fossil occurrence data from 63 well-dated Gondwanan rainforest sites and 396 constituent taxa. Fossil sites were allocated to four age groups: Cretaceous, Paleocene-Eocene, Neogene plus Oligocene, and Pleistocene. We compared the modern and ancient distributions of lineages represented in the fossil record to see if dissimilarity increased with time. We quantified similarity-dissimilarity of composition and taxonomic structure among fossil assemblages, and between fossil and modern assemblages.•, Key Results: Strong similarities between ancient Patagonia and Australia confirmed shared Gondwanan rainforest history, but more of the lineages persisted in Australia. Samples of ancient Australia grouped with the extant floras of Australia, New Guinea, New Caledonia, Fiji, and Mt. Kinabalu. Decreasing similarity through time among the regional floras of Antarctica, Patagonia, New Zealand, and southern Australia reflects multiple extinction events.•, Conclusions: Gondwanan rainforest lineages contribute significantly to modern rainforest community assembly and often co-occur in widely separated assemblages far from their early fossil records. Understanding how and where lineages from ancient Gondwanan assemblages co-occur today has implications for the conservation of global rainforest vegetation, including in the Old World tropics., (© 2014 Botanical Society of America, Inc.)

Premise of the Study: Relationships of leaf size and shape (physiognomy) with climate have been well characterized for woody non-monocotyledonous angiosperms (dicots), allowing the development of models for estimating paleoclimate from fossil leaves. More recently, petiole width of seed plants has been shown to scale closely with leaf mass. By measuring petiole width and leaf area in fossils, leaf mass per area (MA) can be estimated and an approximate leaf life span inferred. However, little is known about these relationships in ferns, a clade with a deep fossil record and with the potential to greatly expand the applicability of these proxies., Methods: We measured the petiole width, MA, and leaf physiognomic characters of 179 fern species from 188 locations across six continents. We applied biomechanical models and assessed the relationship between leaf physiognomy and climate using correlational approaches., Key Results: The scaling relationship between area-normalized petiole width and MA differs between fern fronds and pinnae. The scaling relationship is best modeled as an end-loaded cantilevered beam, which is different from the best-fit biomechanical model for seed plants. Fern leaf physiognomy is not influenced by climatic conditions., Conclusions: The cantilever beam model can be applied to fossil ferns. The lack of sensitivity of leaf physiognomy to climate in ferns argues against their use to reconstruct paleoclimate. Differences in climate sensitivity and biomechanical relationships between ferns and seed plants may be driven by differences in their hydraulic conductivity and/or their differing evolutionary histories of vein architecture and leaf morphology.

The conservation implications of large-scale rainforest clearing and fragmentation on the persistence of functional and taxonomic diversity remain poorly understood. If traits represent adaptive strategies of plant species to particular circumstances, the expectation is that the effect of forest clearing and fragmentation will be affected by species functional traits, particularly those related to dispersal. We used species occurrence data for woody plants in 46 rainforest patches across 75,000 ha largely cleared of forest by the early 1900s to determine the combined effects of area reduction, fragmentation, and patch size on the taxonomic structure and functional diversity of subtropical rainforest. We compiled species trait values for leaf area, seed dry mass, wood density, and maximum height and calculated species niche breadths. Taxonomic structure, trait values (means, ranges), and the functional diversity of assemblages of climbing and free-standing plants in remnant patches were quantified. Larger rainforest patches had higher species richness. Species in smaller patches were taxonomically less related than species in larger patches. Free-standing plants had a high percentage of frugivore dispersed seeds; climbers had a high proportion of small wind-dispersed seeds. Connections between the patchy spatial distribution of free-standing species, larger seed sizes, and dispersal syndrome were weak. Assemblages of free-standing plants in patches showed more taxonomic and spatial structuring than climbing plants. Smaller isolated patches retained relatively high functional diversity and similar taxonomic structure to larger tracts of forest despite lower species richness. The response of woody plants to clearing and fragmentation of subtropical rainforest differed between climbers and slow-growing mature-phase forest trees but not between climbers and pioneer trees. Quantifying taxonomic structure and functional diversity provides an improved basis for conservation planning and management by elucidating the effects of forest-area reduction and fragmentation. Efectos de la Forma de Crecimiento y Atributos Funcionales en la Respuesta de Plantas Leñosas al Desmonte y Fragmentación de Bosque Lluvioso Subtropical., (© 2013 Society for Conservation Biology.)

Background and Aims: A trade-off between shade tolerance and growth in high light is thought to underlie the temporal dynamics of humid forests. On the other hand, it has been suggested that tree species sorting on temperature gradients involves a trade-off between growth rate and cold resistance. Little is known about how these two major trade-offs interact., Methods: Seedlings of Australian tropical and cool-temperate rainforest trees were grown in glasshouse environments to compare growth versus shade-tolerance trade-offs in these two assemblages. Biomass distribution, photosynthetic capacity and vessel diameters were measured in order to examine the functional correlates of species differences in light requirements and growth rate. Species light requirements were assessed by field estimation of the light compensation point for stem growth., Results: Light-demanding and shade-tolerant tropical species differed markedly in relative growth rates (RGR), but this trend was less evident among temperate species. This pattern was paralleled by biomass distribution data: specific leaf area (SLA) and leaf area ratio (LAR) of tropical species were significantly positively correlated with compensation points, but not those of cool-temperate species. The relatively slow growth and small SLA and LAR of Tasmanian light-demanders were associated with narrow vessels and low potential sapwood conductivity., Conclusions: The conservative xylem traits, small LAR and modest RGR of Tasmanian light-demanders are consistent with selection for resistance to freeze-thaw embolism, at the expense of growth rate. Whereas competition for light favours rapid growth in light-demanding trees native to environments with warm, frost-free growing seasons, frost resistance may be an equally important determinant of the fitness of light-demanders in cool-temperate rainforest, as seedlings establishing in large openings are exposed to sub-zero temperatures that can occur throughout most of the year.

Throughout the evolutionary history of plants, drought, shade, and scarcity of nutrients have structured ecosystems and communities globally. Humans have begun to drastically alter the prevalence of these environmental factors with untold consequences for plant communities and ecosystems worldwide. Given limitations in using organ-level traits to predict ecological performance of species, recent advances using tolerances of low resource availability as plant functional traits are revealing the often hidden roles these factors have in structuring communities and are becoming central to classifying plants ecologically. For example, measuring the physiological drought tolerance of plants has increased the predictability of differences among species in their ability to survive drought as well as the distribution of species within and among ecosystems. Quantifying the shade tolerance of species has improved our understanding of local and regional species diversity and how species have sorted within and among regions. As the stresses on ecosystems continue to shift, coordinated studies of whole-plant growth centered on tolerance of low resource availability will be central in predicting future ecosystem functioning and biodiversity. This will require efforts that quantify tolerances for large numbers of species and develop bioinformatic and other techniques for comparing large number of species.

Background and Aims: The contemporary relegation of conifers mainly to cold or infertile sites has been ascribed to low competitive ability, as a result of the hydraulic inefficiency of tracheids and their seedlings' initial dependence on small foliage areas. Here it is hypothesized that, in temperate rainforests, the larger leaves of angiosperms also reduce self-shading and thus enable display of larger effective foliage areas than the numerous small leaves of conifers., Methods: This hypothesis was tested using 3-D modelling of plant architecture and structural equation modelling to compare self-shading and light interception potential of seedlings of six conifers and 12 angiosperm trees from temperate rainforests. The ratio of displayed leaf area to plant mass (LAR(d)) was used to indicate plant light interception potential: LAR(d) is the product of specific leaf area, leaf mass fraction, self-shading and leaf angle., Results: Angiosperm seedlings self-shaded less than conifers, mainly because of differences in leaf number (more than leaf size), and on average their LAR(d) was about twice that of conifers. Although specific leaf area was the most pervasive influence on LAR(d), differences in self-shading also significantly influenced LAR(d) of large seedlings., Conclusions: The ability to deploy foliage in relatively few, large leaves is advantageous in minimizing self-shading and enhancing seedling light interception potential per unit of plant biomass. This study adds significantly to evidence that vegetative traits may be at least as important as reproductive innovations in explaining the success of angiosperms in productive environments where vegetation is structured by light competition.

• Plant light interception efficiency is a crucial determinant of carbon uptake by individual plants and by vegetation. Our aim was to identify whole-plant variables that summarize complex crown architecture, which can be used to predict light interception efficiency. • We gathered the largest database of digitized plants to date (1831 plants of 124 species), and estimated a measure of light interception efficiency with a detailed three-dimensional model. Light interception efficiency was defined as the ratio of the hemispherically averaged displayed to total leaf area. A simple model was developed that uses only two variables, crown density (the ratio of leaf area to total crown surface area) and leaf dispersion (a measure of the degree of aggregation of leaves). • The model explained 85% of variation in the observed light interception efficiency across the digitized plants. Both whole-plant variables varied across species, with differences in leaf dispersion related to leaf size. Within species, light interception efficiency decreased with total leaf number. This was a result of changes in leaf dispersion, while crown density remained constant. • These results provide the basis for a more general understanding of the role of plant architecture in determining the efficiency of light harvesting., (© 2011 The Authors. New Phytologist © 2011 New Phytologist Trust.)

Background and Aims: A long-running debate centres on whether shade tolerance of tree seedlings is mainly a function of traits maximizing net carbon gain in low light, or of traits minimizing carbon loss. To test these alternatives, leaf display, light-interception efficiency, and simulated net daily carbon gain of juvenile temperate evergreens of differing shade tolerance were measured, and how these variables are influenced by ontogeny was queried., Methods: The biomass distribution of juveniles (17-740 mm tall) of seven temperate rainforest evergreens growing in low (approx. 4 %) light in the understorey of a second-growth stand was quantified. Daytime and night-time gas exchange rates of leaves were also determined, and crown architecture was recorded digitally. YPLANT was used to model light interception and carbon gain., Results: An index of species shade tolerance correlated closely with photosynthetic capacities and respiration rates per unit mass of leaves, but only weakly with respiration per unit area. Accumulation of many leaf cohorts by shade-tolerant species meant that their ratios of foliage area to biomass (LAR) decreased more gradually with ontogeny than those of light-demanders, but also increased self-shading; this depressed the foliage silhouette-to-area ratio (STAR), which was used as an index of light-interception efficiency. As a result, displayed leaf area ratio (LAR(d) = LAR × STAR) of large seedlings was not related to species shade tolerance. Self-shading also caused simulated net daily carbon assimilation rates of shade-tolerant species to decrease with ontogeny, leading to a negative correlation of shade tolerance with net daily carbon gain of large (500 mm tall) seedlings in the understorey., Conclusions: The results suggest that efficiency of energy capture is not an important correlate of shade tolerance in temperate rainforest evergreens. Ontogenetic increases in self-shading largely nullify the potential carbon gain advantages expected to result from low respiration rates and long leaf lifespans in shade-tolerant evergreens. The main advantage of their long-lived leaves is probably in reducing the costs of crown maintenance.

• Paleobotanists have long used models based on leaf size and shape to reconstruct paleoclimate. However, most models incorporate a single variable or use traits that are not physiologically or functionally linked to climate, limiting their predictive power. Further, they often underestimate paleotemperature relative to other proxies. • Here we quantify leaf-climate correlations from 92 globally distributed, climatically diverse sites, and explore potential confounding factors. Multiple linear regression models for mean annual temperature (MAT) and mean annual precipitation (MAP) are developed and applied to nine well-studied fossil floras. • We find that leaves in cold climates typically have larger, more numerous teeth, and are more highly dissected. Leaf habit (deciduous vs evergreen), local water availability, and phylogenetic history all affect these relationships. Leaves in wet climates are larger and have fewer, smaller teeth. Our multivariate MAT and MAP models offer moderate improvements in precision over univariate approaches (± 4.0 vs 4.8°C for MAT) and strong improvements in accuracy. For example, our provisional MAT estimates for most North American fossil floras are considerably warmer and in better agreement with independent paleoclimate evidence. • Our study demonstrates that the inclusion of additional leaf traits that are functionally linked to climate improves paleoclimate reconstructions. This work also illustrates the need for better understanding of the impact of phylogeny and leaf habit on leaf-climate relationships., (© 2011 The Authors. New Phytologist © 2011 New Phytologist Trust.)

*When grown in a common light environment, the leaves of shade-tolerant evergreen trees have a larger leaf mass per unit area (LMA) than their light-demanding counterparts, associated with differences in lifespan. Yet plastic responses of LMA run counter to this pattern: shade leaves have smaller LMA than sun leaves, despite often living longer. *We measured LMA and cell wall content, and conducted punch and shear tests, on sun and shade leaves of 13 rainforest evergreens of differing shade tolerance, in order to understand adaptation vs plastic responses of leaf structure and biomechanics to shade. *Species shade tolerance and leaf mechanical properties correlated better with cell wall mass per unit area than with LMA. Growth light environment had less effect on leaf mechanics than on LMA: shade leaves had, on average, 40% lower LMA than sun leaves, but differences in work-to-shear, and especially force-to-punch, were smaller. This was associated with a slightly larger cell wall fraction in shade leaves. *The persistence of shade leaves might reflect unattractiveness to herbivores because they yield smaller benefits (cell contents per area) per unit fracture force than sun leaves. In forest trees, cell wall fraction and force-to-punch are more robust correlates of species light requirements than LMA.

Leaf mass per area (LMA) is one of the most widely measured of all plant functional traits. In deciduous forests, there is similarity between plastic and evolutionary responses of LMA to light gradients. In evergreens, however, LMA is lower in shaded than sunlit individuals of the same species, whereas shade-tolerant evergreens have higher LMA than light-demanders grown under the same conditions. We suggest that this pattern of 'counter-gradient variation' results from some combination of (i) close evolutionary coordination of LMA with leaf lifespan, (ii) selection for different leaf constitutions (relative investment in cell walls versus cell contents) in sun and shade environments and/or (iii) constraints on plasticity as a result of genetic correlations between phenotypes expressed in sun and shade.

Knowledge of leaf chemistry, physiology, and life span is essential for global vegetation modeling, but such data are scarce or lacking for some regions, especially in developing countries. Here we use data from 2021 species at 175 sites around the world from the GLOPNET compilation to show that key physiological traits that are difficult to measure (such as photosynthetic capacity) can be predicted from simple qualitative plant characteristics, climate information, easily measured ("soft") leaf traits, or all of these in combination. The qualitative plant functional type (PFT) attributes examined are phylogeny (angiosperm or gymnosperm), growth form (grass, herb, shrub, or tree), and leaf phenology (deciduous vs. evergreen). These three PFT attributes explain between one-third and two-thirds of the variation in each of five quantitative leaf ecophysiological traits: specific leaf area (SLA), leaf life span, mass-based net photosynthetic capacity (Amass), nitrogen content (N(mass)), and phosphorus content (P(mass)). Alternatively, the combination of four simple, widely available climate metrics (mean annual temperature, mean annual precipitation, mean vapor pressure deficit, and solar irradiance) explain only 5-20% of the variation in those same five leaf traits. Adding the climate metrics to the qualitative PFTs as independent factors in the model increases explanatory power by 3-11% for the five traits. If a single easily measured leaf trait (SLA) is also included in the model along with qualitative plant traits and climate metrics, an additional 5-25% of the variation in the other four other leaf traits is explained, with the models accounting for 62%, 65%, 66%, and 73% of global variation in N(mass), P(mass), A(mass), and leaf life span, respectively. Given the wide availability of the summary climate data and qualitative PFT data used in these analyses, they could be used to explain roughly half of global variation in the less accessible leaf traits (A(mass), leaf life span, N(mass), P(mass)); this can be augmented to two-thirds of all variation if climatic and PFT data are used in combination with the readily measured trait SLA. This shows encouraging possibilities of progress in developing general predictive equations for macro-ecology, global scaling, and global modeling.

It was predicted that relationships of leaf mass per area (LMA) with juvenile shade tolerance will depend on leaf habit, and on whether species are compared at a common age as young seedlings, or at a common size as saplings. A meta-analysis of 47 comparative studies (372 species) was used to test predictions, and the effect of light environment on this relationship. The LMA of evergreens was positively correlated with shade tolerance, irrespective of ontogeny or light environment. The LMA of young seedlings (

Leaf dark respiration (R) is one of the most fundamental physiological processes in plants and is a major component of terrestrial CO2 input to the atmosphere. Still, it is unclear how predictably species vary in R along broad climate gradients. Data for R and other key leaf traits were compiled for 208 woody species from 20 sites around the world. We quantified relationships between R and site climate, and climate-related variation in relationships between R and other leaf traits. Species at higher-irradiance sites had higher mean R at a given leaf N concentration, specific leaf area (SLA), photosynthetic capacity (Amass) or leaf lifespan than species at lower-irradiance sites. Species at lower-rainfall sites had higher mean R at a given SLA or Amass than species at higher-rainfall sites. On average, estimated field rates of R were higher at warmer sites, while no trend with site temperature was seen when R was adjusted to a standard measurement temperature. Our findings should prove useful for modelling plant nutrient and carbon budgets, and for modelling vegetation shifts with climate change.

In Chilean evergreen temperate forest, fern species of the genus Blechnum occur in diverse microhabitats ranging from large gaps to heavily shaded understoreys. We hypothesised that differences in the ecological breadth of three co-occurring Blechnum species would be associated with differences in magnitude of ecophysiological responses to light availability. We quantified the field distribution of each species in relation to diffuse light availability (% canopy openness), and measured in situ variation in photosynthetic capacity (A), dark respiration (R (d)) and specific leaf area (SLA) across the light gradient. The response of SLA of each species was also evaluated in a common garden in two light conditions (understorey and forest edge). The three Blechnum species differed significantly in the range of light environments occupied (breadth: B. chilense > B. hastatum > B. mochaenum). Despite significant interspecific differences in average A and R (d), the response of these traits to light availability did not differ among species. However, there was significant interspecific variation in both the mean value and the plasticity of SLA to light availability, the species with least ecological breadth (B. mochaenum) showing a flatter reaction norm (lower response) than its two congeners. This pattern was also found in the common garden experiment. The adjustment of leaf morphology (SLA) to light availability appears to be an important mechanism of acclimation in these Blechnum species. The narrow range of light environments occupied by B. mochaenum may be at least partly attributable to its inability to display phenotypic plasticity in SLA to changes in light availability.

•  Here we explore the possible role of leaf-level gas exchange traits in determining growth rate differences and competitive interactions between evergreen angiosperms and conifers. •  We compared relationships among photosynthetic capacity (A max ), maximum stomatal conductance (G s ), leaf life span, nitrogen concentration (N) and specific leaf area (SLA), in sun leaves of 23 evergreen angiosperm and 20 conifer populations. •  Despite similar average leaf N mass , conifer leaves lived longer on average (36 months) than angiosperms (25 months). At a standardized leaf N, A mass was higher in angiosperms (56 nmol g -1 s -1 ) than in conifers (36 nmol g -1 s -1 ). Stepwize regression suggested that most of this difference in photosynthetic nitrogen use efficiency could be explained by G s and SLA. Mean G s (on an area basis) of angiosperms was higher than that of conifers (152 vs 117 mmol m 2 s -1 ), but A area -G s relationships were similar for the two groups. At a given leaf N, conifers had lower SLA (projected area basis) than angiosperms. •  Photosynthetic differences probably contribute to the competitive advantage of angiosperm trees over conifers in productive habitats, and may be linked to the greater hydraulic capacity of vessels, enabling angiosperms to develop higher stomatal conductance and therefore sustain higher transpiration rates.

•  The safe site concept could have utility in community ecology if predictive relationships between plant traits and safe site characteristics could be identified. Here we examine the proposal that the nature and relative abundance of safe sites are systematically related to seed size, in an assemblage of 17 woody species in a temperate forest. •  We compared the degree of association of seedlings of each species with elevated microsites, and examined life-history correlates of this variation. •  Seed size explained 45% of interspecific variation in percentage of seedlings growing on logs and other elevated substrates. Neither specific leaf area nor an index of light requirements gave significant increases in explanatory power. Phylogenetically independent contrasts gave a similar, but weaker, relationship with seed size. Most small-seeded species were overrepresented on elevated microsites, whereas two large-seeded taxa were underrepresented on these substrates. Safe sites of small-seeded species were therefore more spatially restricted than those of large-seeded taxa, as elevated surfaces occupied only 8% of the forest floor. •  Safe site differentiation may help to explain the wide range of seed size present within many communities, as well as species coexistence in forests.

Most knowledge of the physiological correlates of interspecific variation in shade tolerance derives from studies of first-year seedlings in artificial environments. The present study relates growth, allocation, foliage turnover, biomass distribution and gas exchange traits to low-light survival of large seedlings (20-100 cm tall) of eight temperate rainforest evergreens under field conditions. Taxa for which natural mortality was not observed in low light during the 14-month study are referred to here as "shade-tolerant" species, and those which did die in the shade are referred to as "light-demanding" species. In low light (2-5% canopy openness), shade-tolerant species had slightly lower light compensation points than light-demanders. Light-demanding species had more plastic aboveground allocation patterns, generally allocating proportionally less aboveground biomass to foliage production than shade-tolerant associates in high light (>10% canopy openness), but more in low light. Foliage turnover was generally much slower in shade-tolerant species (10-40% year -1 ) than in light-demanding species (30-190%). As these differences in leaf retention outweighed variation in allocation, shade-tolerant species displayed higher leaf areas at all light levels. Furthermore, all shade-tolerant species gained leaf area in low light during the study period, whereas light-demanding taxa showed leaf area declines. Higher leaf area ratios, plus differences in light compensation points, indicate that large seedlings of shade-tolerant evergreens enjoy net carbon gain advantages over light-demanding associates in low light. However, minimal growth rate differences in low light imply higher storage allocation in shade-tolerant species. This study provides a rather different picture from that which has emerged from recent reviews of first-year seedling data, illustrating the long-term consequences of foliage turnover differences for biomass distribution, and suggesting that shade tolerance in juvenile evergreen trees is associated with a suite of traits which enhance net carbon gain, but not growth, in low light. Accumulation of a large foliage area through long leaf retention times is probably a key mechanism enhancing low-light carbon gain in evergreens.

•  In a comparative study of 26 Chilean temperate woody species, the following questions were addressed. Are germination requirements correlated with seedling light requirements? Can germination requirements be used to delimit pioneer and nonpioneer guilds? •  To assess dependence of germination on canopy gaps, germination percentages were compared experimentally in gap and understorey environments. Seedling light requirements were quantified by determining the mean light environment naturally occupied by juveniles of each species. •  Germination was significantly higher in the gap than in the understorey in > 30% of the species, although only one germinated exclusively in the gap. In the understorey, germination was higher in 19% of species, whereas 50% were indifferent to germination environment. Several light-demanding species germinated abundantly in the understorey. Both cross-species analysis and phylogenetically independent contrasts showed that seedling light requirements were not correlated with responsiveness of germination to the gap environment. •  Results suggest that germination requirements and seedling light requirements are not coupled, and that requirement of canopy gaps for germination cannot be used to delimit pioneer and nonpioneer guilds in a Chilean temperate rain forest.

It has been argued that plants adapted to low light should have lower carbon losses via dark respiration (Rd) than those not so adapted, and similarly, all species would be expected to down-regulate Rd in deep shade, because the associated advantages of high metabolic potential cannot be realized in such habitats. In order to test these hypotheses, and to explore the determinants of intraspecific variation in respiration rates, we measured Rd, leaf mass per unit area (LMA), and nitrogen content of mature foliage in juveniles of 11 cold-temperate tree species (angiosperms and conifers), growing in diverse light environments in forest understories in northern Minnesota. Among the seven angiosperm species, respiration on mass, area, and nitrogen bases showed significant negative overall relationships with shade tolerance level. Mass-based respiration rates (Rd mass ) of angiosperms as a group showed a significant positive overall relationship with an index of light availability (percentage canopy openness, %CO). Rd mass of most conifers also showed evidence of acclimation of Rd mass to light availability. LMA of all species also increased with increasing %CO, but this response was generally much stronger in angiosperms than in conifers. As a result, the response of area-based respiration (Rd area ) to %CO was dominated by ΔRd mass for conifers, and by ΔLMA for most angiosperms, i.e., functional types differed in the components of acclimation of Rd area to light availability. Among the seven angiosperm species, the relationships of leaf N on a mass basis (N mass ) with %CO were modulated by shade tolerance: negative slopes in shade-tolerant species may be related to the steep increases in LMA of these taxa along gradients of increasing light intensity, and associated dilution of N-rich, metabolically active tissue by increasing investment in leaf structural components. Although N mass was therefore an unreliable predictor of variation in Rd mass along light gradients, respiration per unit leaf N (Rd/N) was significantly positively correlated with %CO for most species. This probably reflects variation in the proportion of leaf N allocated to protein and/or the influence of leaf carbohydrate status on Rd. Species shade tolerance differences were not significantly correlated with the magnitude of either ΔRd mass or ΔRd area , indicating that variation in acclimation potential of Rd is much less important than inherent differences in this trait. Acclimation of Rd mass to light availability appears to be a generalized feature of juvenile trees, and the important ecological trade-off is likely between high metabolic capacity in high light and low respiratory losses in low light.

Seedlings of nine southern Chilean trees were grown at three nutrient supply rates, to examine the roles of growth rate, biomass distribution and nutrient use traits in determining species natural distributions on resource gradients. Relative growth rate (RGR) showed no overall relationship with species site requirements, although RGR of fertile-site species tended to be more responsive to nutrient supply. In the low-nutrient treatment, infertility-tolerant Fitzroya cupressoides showed a higher RGR rank than a fertility-demanding species (Laurelia philippiana) which outgrew it substantially at the highest supply rate. This reversal of RGR ranks was associated with divergent nutrient use responses: at high nutrient supply both spp. had similar plant nitrogen concentrations (PNC), whereas at the low supply rate Fitzroya's production of biomass per unit of assimilated N was twice that of Laurelia's. However, this pattern does not appear to serve as a general explanation of the respective distributions of the study species, as RGR ranks of most species were unaltered by nutrient supply. At low nutrient availability, no clear differences in shoot:root ratio (SRR) were apparent between poor-site and fertile-site species. However, at high nutrient availability, SRR was markedly higher in the latter, resulting from differences in biomass allocation to stems (not leaves). Leaf area ratios (LAR) were higher in fertile-site species than in those tolerant of low fertility, because of differences in specific leaf area rather than leaf weight ratio. Very high LAR at high nutrient supply was characteristic of most shade-tolerant angiosperms, but not of shade-tolerant conifers. Although PNC showed no overall differences between poor- and fertile-site species, sensitivity of PNC to external supply rate was greatest in two infertility-tolerant conifers. In contrast, the angiosperm Weinmannia trichosperma, although tolerant of low fertility, responded to increased nutrient supply with greatly increased RGR and little change in PNC. Results show little trait convergence between conifers and angiosperms in adaptation both to shade and to infertile soils; i.e. fitness of different taxa in a given environment may hinge on different trait combinations.