U.S. National Arboretum

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives.

ABSTRACT The hydraulic conductance of the leaf lamina ( K lamina ) substantially constrains whole‐plant water transport, but little is known of its association with leaf structure and function. K lamina was measured for sun and shade leaves of six woody temperate species growing in moist soil, and tested for correlation with the prevailing leaf irradiance, and with 22 other leaf traits. K lamina varied from 7.40 × 10 −5 kg m −2 s −1 MPa −1 for Acer saccharum shade leaves to 2.89 × 10 −4 kg m −2 s −1 MPa −1 for Vitis labrusca sun leaves. Tree sun leaves had 15–67% higher K lamina than shade leaves. K lamina was co‐ordinated with traits associated with high water flux, including leaf irradiance, petiole hydraulic conductance, guard cell length, and stomatal pore area per lamina area. K lamina was also co‐ordinated with lamina thickness, water storage capacitance, 1/mesophyll water transfer resistance, and, in five of the six species, with lamina perimeter/area. However, for the six species, K lamina was independent of inter‐related leaf traits including leaf dry mass per area, density, modulus of elasticity, osmotic potential, and cuticular conductance. K lamina was thus co‐ordinated with structural and functional traits relating to liquid‐phase water transport and to maximum rates of gas exchange, but independent of other traits relating to drought tolerance and to aspects of carbon economy.

BACKGROUND: The characteristics of pollen tube growth are not constant, but display distinct patterns of growth within the different tissues of the pistil. In the stigma, the growth rate is slow and autotrophic, whereas in the style, it is rapid and heterotrophic. Very little is known about the interactions between these distinct maternal tissues and the traversing pollen tube and the role of this interaction on the observed metabolism. In this work we characterise pollen tube growth in the apple flower and look for differences in glycoprotein epitope localization between two different maternal tissues, the stigma and the style. RESULTS: While immunocytochemically-detected arabinogalactan proteins were present at high levels in the stigma, they were not detected in the transmitting tissue of the style, where extensins were abundant. Whereas extensins remained at high levels in unpollinated pistils, they were no longer present in the style following pollen tube passage. Similarily, while abundant in unpollinated styles, insoluble polysaccharides such as β-glucans, were depleted in pollinated pistils. CONCLUSIONS: The switch from autotropic to heterotrophic pollen tube growth correlates spatially with a change of glycoprotein epitopes between the stigma and the style. The depletion of extensins and polysaccharides following pollen tube passage in the style suggest a possible contribution to the acceleration of heterotrophic pollen tube growth, which would imply an active contribution of female tissues on prezygotic male-female crosstalk.

19:2348-2359], suggesting that shifts toward asynchrony may be infrequent. A meta-analytic approach would provide insights into global trends and how they are linked to climate change. We compared phenological shifts among pairwise species interactions (e.g., predator-prey) using published long-term time-series data of phenological events from aquatic and terrestrial ecosystems across four continents since 1951 to determine whether recent climate change has led to overall shifts in synchrony. We show that the relative timing of key life cycle events of interacting species has changed significantly over the past 35 years. Further, by comparing the period before major climate change (pre-1980s) and after, we show that estimated changes in phenology and synchrony are greater in recent decades. However, there has been no consistent trend in the direction of these changes. Our findings show that there have been shifts in the timing of interacting species in recent decades; the next challenges are to improve our ability to predict the direction of change and understand the full consequences for communities and ecosystems.

The veins that irrigate leaves during photosynthesis are demonstrated to be strikingly more abundant in flowering plants than in any other vascular plant lineage. Angiosperm vein densities average 8 mm of vein per mm(2) of leaf area and can reach 25 mm mm(-2), whereas such high densities are absent from all other plants, living or extinct. Leaves of non-angiosperms have consistently averaged close to 2 mm mm(-2) throughout 380 million years of evolution despite a complex history that has involved four or more independent origins of laminate leaves with many veins and dramatic changes in climate and atmospheric composition. We further demonstrate that the high leaf vein densities unique to the angiosperms enable unparalleled transpiration rates, extending previous work indicating a strong correlation between vein density and assimilation rates. Because vein density is directly measurable in fossils, these correlations provide new access to the physiology of extinct plants and how they may have impacted their environments. First, the high assimilation rates currently confined to the angiosperms among living plants are likely to have been unique throughout evolutionary history. Second, the transpiration-driven recycling of water that is important for bolstering precipitation in modern tropical rainforests might have been significantly less in a world before the angiosperms.

Summary New analytical tools applied to long‐term data demonstrate that ecological communities are highly dynamic over time. We developed an r package, library(“codyn”) , to help ecologists easily implement these metrics and gain broader insights into ecological community dynamics. library(“codyn”) provides temporal diversity indices and community stability metrics. All functions are designed to be easily implemented over multiple replicates. Temporal diversity indices include species turnover, mean rank shifts and rate of community change over time. Community stability metrics calculate overall stability and patterns of species covariance and synchrony over time, and include a null‐modelling method to test significance. Finally, library(“codyn”) contains vignettes that describe methods and reproduce figures from published papers to help users contextualize and apply functions to their own data.

Two fundamental axes - space and time - shape ecological systems. Over the last 30 years spatial ecology has developed as an integrative, multidisciplinary science that has improved our understanding of the ecological consequences of habitat fragmentation and loss. We argue that accelerating climate change - the effective manipulation of time by humans - has generated a current need to build an equivalent framework for temporal ecology. Climate change has at once pressed ecologists to understand and predict ecological dynamics in non-stationary environments, while also challenged fundamental assumptions of many concepts, models and approaches. However, similarities between space and time, especially related issues of scaling, provide an outline for improving ecological models and forecasting of temporal dynamics, while the unique attributes of time, particularly its emphasis on events and its singular direction, highlight where new approaches are needed. We emphasise how a renewed, interdisciplinary focus on time would coalesce related concepts, help develop new theories and methods and guide further data collection. The next challenge will be to unite predictive frameworks from spatial and temporal ecology to build robust forecasts of when and where environmental change will pose the largest threats to species and ecosystems, as well as identifying the best opportunities for conservation.

Summary Over the past several years, phylogenetic comparative studies have increasingly approached trait evolution in a multivariate context, with a number of taxa that continues to rise dramatically. Recent methods for phylogenetic comparative studies have provided ways to incorporate measurement error and to address computational challenges. However, missing data remain a particularly common problem, in which data are unavailable for some but not all traits of interest for a given species (or individual), leaving researchers with the choice between omitting observations or utilizing imputation‐based approaches. Here, we introduce an r implementation of PhyloPars , a tool for phylogenetic imputation of missing data and estimation of trait covariance across species (phylogenetic covariance) and within species (phenotypic covariance). Rphylopars provides expanded capabilities over the original PhyloPars interface including a fast linear‐time algorithm, thus allowing for extremely large data sets (which were previously computationally infeasible) to be analysed in seconds or minutes rather than hours. In addition to providing fast and computationally efficient implementations, we introduce in Rphylopars methods to estimate macroevolutionary parameters under alternative evolutionary models (e.g. Early‐Burst, multivariate Ornstein‐Uhlenbeck). By providing fast and computationally efficient methods with flexible options for various phylogenetic comparative approaches, Rphylopars expands the possibilities for researchers to analyse large and complex data with missing observations, within‐species variation and deviations from Brownian motion.

Geological and climatological processes that have impacted the biota of the Northern Hemisphere during the Tertiary are expected to yield little resolution when area cladograms are compared without taking the timing of diversification into account. In an attempt to establish a set of appropriate phylogenetic comparisons, we distinguished between a Pacific track involving (minimally) China, Japan, and eastern North America but not Europe, and an Atlantic track involving China, Europe, and eastern North America but not Japan (or, in most cases, western North America). Within the two Atlantic‐track taxa considered here—Liquidambar and Cercis—European and North American species are more closely related to one another than they are to the Asian species. Within a set of five Pacific‐track taxa—Hamamelis, Weigela‐Diervilla, Triosteum, Buckleya, and Torreya—we see all possible relationships involving China, Japan, and eastern North America. Estimates of minimum divergence times between Old World and New World lineages, based on molecular and fossil evidence, differ markedly between the two Atlantic‐track clades. Among the Pacific‐track taxa, we find no correlation between pattern of area relationships and estimated divergence times of the Old World–New World disjuncts. Instead, we see a wide range in the timing of these splitting events among and within phylogenetic patterns. Despite the existence of a variety of patterns, inferred ancestral areas and divergence times can be explained by assuming initial diversification within Asia in a number of lineages, followed by iterative trans‐Beringian dispersion and vicariance.

How climate affects species distributions is a longstanding question receiving renewed interest owing to the need to predict the impacts of global warming on biodiversity. Is climate change forcing species to live near their critical thermal limits? Are these limits likely to change through natural selection? These and other important questions can be addressed with models relating geographical distributions of species with climate data, but inferences made with these models are highly contingent on non-climatic factors such as biotic interactions. Improved understanding of climate change effects on species will require extensive analysis of thermal physiological traits, but such data are both scarce and scattered. To overcome current limitations, we created the GlobTherm database. The database contains experimentally derived species' thermal tolerance data currently comprising over 2,000 species of terrestrial, freshwater, intertidal and marine multicellular algae, plants, fungi, and animals. The GlobTherm database will be maintained and curated by iDiv with the aim to keep expanding it, and enable further investigations on the effects of climate on the distribution of life on Earth.

Mast-fruiting Dipterocarpaceae exhibit highly synchronous, interspecific seedfall at irregular, multiyear intervals. To investigate how the temporal pattern of seedfall affects dipterocarp seed and seedling survival, in both a logged and a primary lowland tropical forest, we planted Shorea stenoptera Burck seeds in the last three weeks of a 12-wk synchronous dipterocarp seedfall during a major community mast-fruiting event in West Kalimantan, Indonesia. As a result of commercial timber harvest of dipterocarp individuals eight years before, total dipterocarp seed production in the logged site was only 23% of that in the primary forest. At both sites, an average of 35 kg of seed was sown across a large area (≥1 km2) to examine the spatial pattern of seed destruction. During the period in which “natural” community mast seed was available to predators, 92% and 99% of experimentally sown seed escaped predators in logged and in primary forest, respectively. After regional seed resources were exhausted, nomadic vertebrates (primarily the bearded pig, Sus barbatus) were observed in both forest areas, and all ungerminated seed was destroyed. Seed predators arrived earlier in the logged area, before most experimentally sown seed had germinated, and the logged site experienced greater seed loss to vertebrates than did the primary forest. Because nomadic seed predators were absent during peak fruit fall of naturally occurring communities at both study sites, there was no evidence of local predator satiation. Rather, experimentally sown seed escaped predation because of rapid germination before predator arrival, as opposed to being ignored by satiated predators. Seed escape was more dependent on the late arrival of pigs than on the amount of local seed production. There was no significant spatial autocorrelation of seed predation. All remaining seed at the scale of the experiment (>1 km2) was destroyed by predators. These findings suggest that satiation of nomadic predators occurs at the landscape scale. Postdispersal seed predators caused significantly greater seed destruction in the experimentally sown seed populations than in naturally dispersed, mast-fruiting communities at both sites. In both logged and primary forests, there was significantly greater loss of experimentally sown seed to predation than was found in the entire natural mast-fruiting Shorea community combined (21 spp.). Moreover, a naturally occurring, but late-fruiting, Shorea species also exhibited greater seed losses to predation than did all other species within each mast-fruiting community, and these proportional losses were similar to those observed in the experimentally sown seeds. Seeds that escaped predation and vertebrate herbivory on post-establishment seedlings displayed high survival, indicating that the availability of suitable microsites did not limit recruitment. In the primary forest, 65% of the germinated experimental seed that survived early causes of mortality was alive 40 mo post-planting, which coincided with the next mast-fruiting event. The spatial distribution of these seedlings was modified primarily by the foraging behavior of vertebrate seed predators in the first two weeks post-planting. The influence of vertebrate predation on seed and seedling survival suggests that foraging behavior by terrestrial vertebrate seed predators may cause directional and/or stabilizing selection for synchronous, interspecific supra-annual dipterocarp seed production across forest regions in Kalimantan.

The family Brassicaceae comprises 3710 species in 338 genera, 25 recently delimited tribes, and three major lineages based on phylogenetic results from the chloroplast gene ndhF. To assess the credibility of the lineages and newly delimited tribes, we sequenced an approximately 1.8-kb region of the nuclear phytochrome A (PHYA) gene for taxa previously sampled for the chloroplast gene ndhF. Using parsimony, likelihood, and Bayesian methods, we reconstructed the phylogeny of the gene and used the approximately unbiased (AU) test to compare phylogenetic results from PHYA with findings from ndhF. We also combined ndhF and PHYA data and used a Bayesian mixed model approach to infer phylogeny. PHYA and combined analyses recovered the same three large lineages as those recovered in ndhF trees, increasing confidence in these lineages. The combined tree confirms the monophyly of most of the recently delimited tribes (only Alysseae, Anchonieae, and Descurainieae are not monophyletic), while 13 of the 23 sampled tribes are monophyletic in PHYA trees. In addition to phylogenetic results, we documented the trichome branching morphology of species across the phylogeny and explored the evolution of different trichome morphologies using the AU test. Our results indicate that dendritic, medifixed, and stellate trichomes likely evolved independently several times in the Brassicaceae.

Agrobiodiversity—the variation within agricultural plants, animals, and practices—is often suggested as a way to mitigate the negative impacts of climate change on crops [S. A. Wood et al. , Trends Ecol. Evol. 30, 531–539 (2015)]. Recently, increasing research and attention has focused on exploiting the intraspecific genetic variation within a crop [Hajjar et al. , Agric. Ecosyst. Environ. 123, 261–270 (2008)], despite few relevant tests of how this diversity modifies agricultural forecasts. Here, we quantify how intraspecific diversity, via cultivars, changes global projections of growing areas. We focus on a crop that spans diverse climates, has the necessary records, and is clearly impacted by climate change: winegrapes (predominantly Vitis vinifera subspecies vinifera ). We draw on long-term French records to extrapolate globally for 11 cultivars (varieties) with high diversity in a key trait for climate change adaptation—phenology. We compared scenarios where growers shift to more climatically suitable cultivars as the climate warms or do not change cultivars. We find that cultivar diversity more than halved projected losses of current winegrowing areas under a 2 °C warming scenario, decreasing areas lost from 56 to 24%. These benefits are more muted at higher warming scenarios, reducing areas lost by a third at 4 °C (85% versus 58%). Our results support the potential of in situ shifting of cultivars to adapt agriculture to climate change—including in major winegrowing regions—as long as efforts to avoid higher warming scenarios are successful.

• The leaf hydraulic conductance (Kleaf) is a major determinant of plant water transport capacity. Here, we measured Kleaf, and its basis in the resistances of leaf components, for fully illuminated leaves of five tree species that regenerate in deep shade, and five that regenerate in gaps or clearings, in Panamanian lowland tropical rainforest. We also determined coordination with stomatal characters and leaf mass per area. • K leaf varied 10-fold across species, and was 3-fold higher in sun- than in shade-establishing species. On average, 12% of leaf hydraulic resistance (= 1/Kleaf) was located in the petiole, 25% in the major veins, 25% in the minor veins, and 39% outside the xylem. Sun-establishing species had a higher proportion of leaf resistance in the xylem. Across species, component resistances correlated linearly with total leaf resistance. • K leaf correlated tightly with indices of stomatal pore area, indicating a coordination of liquid- and vapor-phase conductances shifted relative to that of temperate woody species. • Leaf hydraulic properties are integrally linked in the complex of traits that define differences in water use and carbon economy across habitats and vegetation zones.

Partial sequences of the nuclear gene encoding the photoreceptor phytochrome A (PHYA) are used to reconstruct relationships within Orobanchaceae, the largest of the parasitic angiosperm families. The monophyly of Orobanchaceae, including nonphotosynthetic holoparasites, hemiparasites, and nonparasitic Lindenbergia is strongly supported. Phytochrome A data resolve six well-supported lineages that contain all of the sampled genera except Brandisia, which is sister to the major radiation of hemiparasites. In contrast to previous plastid and ITS trees, relationships among these major clades also are generally well supported. Thus, the robust phylogenetic hypothesis inferred from the PHYA data provides a much better context in which to evaluate the evolution of parasitism within the group. Ninety-eight species of Orobanchaceae, representing 43 genera, are included and Brandisia, Bungea, Cymbaria, Esterhazya, Nesogenes, Phtheirospermum, Radamaea, Siphonostegia, and Xylocalyx are confirmed as members of Orobanchaceae. The earliest diverging lineage of hemiparasites is identified for the first time; it contains Bungea, Cymbaria, Monochasma, Siphonostegia, and the monotypic Schwalbea, which is federally endangered. This basal clade is marked by the presence of two novel introns. A second, apparently independent gain of one of these introns marks a clade of largely European taxa. There is significant rate heterogeneity among PHYA sequences, and the presence of multiple PHYA in some taxa is consistent with observed ploidy levels.

Leaves constitute a substantial fraction of the total resistance to water flow through plants. A key question is how hydraulic resistance within the leaf is distributed among petiole, major veins, minor veins, and the pathways downstream of the veins. We partitioned the leaf hydraulic resistance (R(leaf)) for sugar maple (Acer saccharum) and red oak (Quercus rubra) by measuring the resistance to water flow through leaves before and after cutting specific vein orders. Simulations using an electronic circuit analog with resistors arranged in a hierarchical reticulate network justified the partitioning of total R(leaf) into component additive resistances. On average 64% and 74% of the R(leaf) was situated within the leaf xylem for sugar maple and red oak, respectively. Substantial resistance-32% and 49%- was in the minor venation, 18% and 21% in the major venation, and 14% and 4% in the petiole. The large number of parallel paths (i.e. a large transfer surface) for water leaving the minor veins through the bundle sheath and out of the leaf resulted in the pathways outside the venation comprising only 36% and 26% of R(leaf). Changing leaf temperature during measurement of R(leaf) for intact leaves resulted in a temperature response beyond that expected from changes in viscosity. The extra response was not found for leaves with veins cut, indicating that water crosses cell membranes after it leaves the xylem. The large proportion of resistance in the venation can explain why stomata respond to leaf xylem damage and cavitation. The hydraulic importance of the leaf vein system suggests that the diversity of vein system architectures observed in angiosperms may reflect variation in whole-leaf hydraulic capacity.

Intracanopy plasticity in tree leaf form is a major determinant of whole-plant function and potentially of forest understory ecology. However, there exists little systematic information for the full extent of intracanopy plasticity, whether it is linked with height and exposure, or its variation across species. For arboretum-grown trees of six temperate deciduous species averaging 13-18 m in height, we quantified intracanopy plasticity for 11 leaf traits across three canopy locations (basal-interior, basal-exterior, and top). Plasticity was pronounced across the canopy, and maximum likelihood analyses indicated that plasticity was primarily linked with irradiance, regardless of height. Intracanopy plasticity (the quotient of values for top and basal-interior leaves) was often similar across species and statistically indistinguishable across species for several key traits. At canopy tops, the area of individual leaves was on average 0.5-0.6 times that at basal-interior, stomatal density 1.1-1.5 times higher, sapwood cross-sectional area up to 1.7 times higher, and leaf mass per area 1.5-2.2 times higher; guard cell and stomatal pore lengths were invariant across the canopy. Species differed in intracanopy plasticity for the mass of individual leaves, leaf margin dissection, ratio of leaf to sapwood areas, and stomatal pore area per leaf area; plasticity quotients ranged only up to ≈2. Across the six species, trait plasticities were uncorrelated and independent of the magnitude of the canopy gradient in irradiance or height and of the species' light requirements for regeneration. This convergence across species indicates general optimization or constraints in development, resulting in a bounded plasticity that improves canopy performance.

Abstract An intensive soil sampling design was evaluated to determine what resolution could be obtained in N and C pool size estimates in a northern hardwood forest soil. Pits of measured volume were excavated by horizon in the forest floor and in three depth strata in the mineral soil. Future comparisons should be able to detect differences in N and C pool sizes ranging from 8 to 25% of the observed mean values depending upon the element and depth strata. Future sampling should detect changes of 230 and 130 kg N ha −1 in the forest floor (combined O horizons) and 0‐ to 10‐cm stratum in the mineral soil respectively. Similarly, changes of 5.9 and 2.4 Mg C ha −1 should be detectable for forest floor and 0 to 10 cm pools respectively. Soil N content for the forest floor was 1300 kg N ha −1 . For the mineral soil depth strata (0–10 cm, 10–20 cm, 20 cm to the bottom of the B horizon), N contents were 1600, 1200 and 3100 kg N ha −1 respectively. Total solum N content was estimated to be 7200 kg N ha −1 . Soil C contents for the combined O horizons, 0‐ to 10‐, 10‐ to 20‐ and ≥ 20‐cm strata were 30, 32, 27 and 73 Mg C ha −1 respectively. The total solum C content was estimated to be 160 Mg C ha −1 . Concentrations of soil N and C were positively correlated with elevation over the 240 m range studied, but soil pools of N and C were not correlated with elevation or soil mapping unit.

The long evolution of vascular plants has resulted in a tremendous variety of natural networks responsible for the evaporatively driven transport of water. Nevertheless, little is known about the physical principles that constrain vascular architecture. Inspired by plant leaves, we used microfluidic devices consisting of simple parallel channel networks in a polymeric material layer, permeable to water, to study the mechanisms of and the limits to evaporation-driven flow. We show that the flow rate through our biomimetic leaves increases linearly with channel density (1/d) until the distance between channels (d) is comparable with the thickness of the polymer layer (delta), above which the flow rate saturates. A comparison with the plant vascular networks shows that the same optimization criterion can be used to describe the placement of veins in leaves. These scaling relations for evaporatively driven flow through simple networks reveal basic design principles for the engineering of evaporation-permeation-driven devices, and highlight the role of physical constraints on the biological design of leaves.

Soil ecologists have debated the relative importance of dispersal limitation and ecological factors in determining the structure of soil microbial communities. Recent evidence suggests that 'everything is not everywhere', and that microbial communities are influenced by both dispersal limitation and ecological factors. However, we still do not understand the relative explanatory power of spatial and ecological factors, including plant species identity and even plant relatedness, for different fractions of the soil microbial community (i.e. bacterial and fungal communities). To ask whether factors such as plant species, soil chemistry, spatial location and plant relatedness influence rhizosphere community composition, we examined field-collected rhizosphere soil of seven congener pairs that occur at Bodega Bay Marine Reserve, CA, USA. We characterized differences in bacterial and fungal communities using terminal-restriction fragment length polymorphism. Plant species identity was the single best statistical predictor of both bacterial and fungal community composition in the root zone. Soil microbial community structure was also correlated with soil chemistry. The third best predictor of bacterial and fungal communities was spatial location, confirming that everything is not everywhere. Variation in microbial community composition was also related to combinations of spatial location, soil chemistry and plant relatedness, suggesting that these factors do not act independently. Plant relatedness explained less of the variation than plant species, soil chemistry, or spatial location. Despite some congeners occupying different habitats and being spatially distant, rhizosphere fungal communities of plant congeners were more similar than expected by chance. Bacterial communities from the same samples were only weakly similar between plant congeners. Thus, plant relatedness might influence soil fungal, more than soil bacterial, community composition.