Comparative venation costs of monocotyledon and dicotyledon species in the eastern Colorado steppe.

MAIN CONCLUSION: Leaf vein network cost (total vein surface area per leaf volume) for major veins and vascular bundles did not differ between monocot and dicot species in 21 species from the eastern Colorado steppe. Dicots possessed significantly larger minor vein networks than monocots. Across the tree of life, there is evidence that dendritic vascular transport networks are optimized, balancing maximum speed and integrity of resource delivery with minimal resource investment in transport and infrastructure. Monocot venation, however, is not dendritic, and remains parallel down to the smallest vein orders with no space-filling capillary networks. Given this departure from the "optimized" dendritic network, one would assume that monocots are operating at a significant energetic disadvantage. In this study, we investigate whether monocot venation networks bear significantly greater carbon/construction costs per leaf volume than co-occurring dicots in the same ecosystem, and if so, what physiological or ecological advantage the monocot life form possesses to compensate for this deficit. Given that venation networks could also be optimized for leaf mechanical support or provide herbivory defense, we measured the vascular system of both monocot and dicots at three scales to distinguish between leaf investment in mechanical support (macroscopic vein), total transport and capacitance (vascular bundle), or exclusively water transport (xylem) for both parallel and dendritic venation networks. We observed that vein network cost (total vein surface area per leaf volume) for major veins and vascular bundles was not significantly different between monocot species and dicot species. Dicots, however, possess significantly larger minor vein networks than monocots. The 19 species subjected to gas-exchange measurement in the field displayed a broad range of Amax and but demonstrated no significant relationships with any metric of vascular network size in major or minor vein classes. Given that monocots do not seem to display any leaf hydraulic disadvantage relative to dicots, it remains an important research question why parallel venation (truly parallel, down to the smallest vessels) has not arisen more than once in the history of plant evolution.

Searching for mechanisms driving root pressure in Zea mays-a transcriptomic approach.

While there are many theories and a variety of innovative datasets contributing to our understanding of the mechanism generating root pressure in vascular plants, we are still unable to produce a specific cellular mechanism for any species. To discover these mechanisms, we used RNA-Seq to explore differentially expressed genes in three different tissues between individual Zea mays plants expressing root pressure and those producing none. Working from the perspective that roots cells are utililizing a combination of osmotic exudation and hydraulic pressure mechanisms to generate positively-pressured flow of water into the xylem from the soil, we hypothesized that differential expression analysis would yield candidate genes coding for membrane transporters, ion channels, ATPases, and hormones with clear relevance to root pressure generation. In basal stem and coarse root tissue, we observed these classes of differentially expressed genes and more, including a strong cytoskeletal remodeling response. Fine roots displayed remarkably little differential expression relevant to root pressure, leading us to conclude that they either do not contribute to root pressure generation or are constitutively expressing root pressure mechanisms regardless of soil water content.

Development and application of an inexpensive open-source dendrometer for detecting xylem water potential and radial stem growth at high spatial and temporal resolution.

There is currently a need for inexpensive, continuous, non-destructive water potential measurements at high temporal resolution (<1 min). We describe here the development and testing of an entirely open-source dendrometer that, when combined with periodic Scholander pressure chamber measurements, provides sub-minute resolution estimates of water potential when placed on tissues exhibiting little or no secondary growth (petioles, monocotyledon stems). The dendrometer can also be used to measure radial growth of stems and branches when placed on dicotyledon and gymnosperm species. The dendrometer can be interfaced directly with a computer in real time in the lab or greenhouse, or connected to a datalogger for long periods of use in the field on batteries. We tested this device on a herbaceous dicotyledon (Helianthus annuus) (petioles and stems) and a monocotyledon (Zea mays) species (stems) for 1 week during dehydration and re-watering treatments under laboratory conditions. We also demonstrated the ability of the device to record branch and trunk diameter variation of a woody dicotyledon (Rhus typhina) in the field. Under laboratory conditions, we compared our device (hereafter 'contact' dendrometer) with modified versions of another open-source dendrometer (the 'optical' dendrometer). Overall, contact and optical dendrometers were well aligned with one another, with Pearson correlation coefficients ranging from 0.77 to 0.97. Both dendrometer devices were well aligned with direct measurements of xylem water potential, with calibration curves exhibiting significant non-linearity, especially at water potentials near the point of incipient plasmolysis, with pseudo R2 values (Efron) ranging from 0.89 to 0.99. Overall, both dendrometers were comparable and provided sufficient resolution to detect subtle differences in stem water potential (ca. 50 kPa) resulting from light-induced changes in transpiration, vapour pressure deficit and drying/wetting soils. All hardware designs, alternative configurations, software and build instructions for the contact dendrometers are provided.

Grass veins are leaky pipes: vessel widening in grass leaves explain variation in stomatal conductance and vessel diameter among species.

The widening of xylem vessels from tip to base of trees is an adaptation to minimize the hydraulic resistance of a long pathway. Given that parallel veins of monocot leaves do not branch hierarchically, vessels should also widen basipetally but, in addition to minimizing resistance, should also account for water volume lost to transpiration since they supply water to the lamina along their lengths, that is 'leakiness'. We measured photosynthesis, stomatal conductance, and vessel diameter at five locations along each leaf of five perennial grass species. We found that the rate of conduit widening in grass leaves was larger than the widening exponent required to minimize pathlength resistance (0.35 vs c. 0.22). Furthermore, variation in the widening exponent among species was positively correlated with maximal stomatal conductance (r2 = 0.20) and net CO2 assimilation (r2 = 0.45). These results suggest that faster rates of conduit widening (> 0.22) were associated with higher rates of water loss. Taken together, our results show that the widening exponent is linked to plant function in grass leaves and that natural selection has favored parallel vein networks that are constructed to meet transpiration requirements while minimizing hydraulic resistance within grass blades.

The effects of copper deficiency on lignification, xylem vessel structure, and hydraulic traits in hybrid poplar.

Copper (Cu) homeostasis is integral to many plant physiological processes, including lignification of plant cell walls. This link occurs through Cu's role as a cofactor in the apoplastic laccase enzymes that oxidize monolignols that then polymerize to form the hydrophobic lignin polymer, which provides rigidity and strength to the water transport system. In this study, we investigated the effect of Cu deficiency on lignin content and chemistry in poplar stems. We also examined the effect of Cu deficiency on the stiffness of stem wood and the hydraulic properties of leaves. Cu deficiency resulted in a significant reduction in lignin content, an increase in the syringyl to guaiacyl monomer ratio of stem xylem, and no change to stem modulus of elasticity. Accompanying these stem traits, Cu-deficient leaves had higher (less negative) turgor loss points and markedly stiffer mesophyll cell walls. Our results may reflect a novel response in poplar whereby structural stiffness and mechanical stability are maintained in the face of Cu deficiency and reduction in the guaiacyl lignin monomer content.

Soil-plant hydraulics explain stomatal efficiency-safety tradeoff.

The efficiency-safety tradeoff has been thoroughly investigated in plants, especially concerning their capacity to transport water and avoid embolism. Stomatal regulation is a vital plant behaviour to respond to soil and atmospheric water limitation. Recently, a stomatal efficiency-safety tradeoff was reported where plants with higher maximum stomatal conductance (gmax ) exhibited greater sensitivity to stomatal closure during soil drying, that is, less negative leaf water potential at 50% gmax (ψgs50 ). However, the underlying mechanism of this gmax -ψgs50 tradeoff remains unknown. Here, we utilized a soil-plant hydraulic model, in which stomatal closure is triggered by nonlinearity in soil-plant hydraulics, to investigate such tradeoff. Our simulations show that increasing gmax is aligned with less negative ψgs50 . Plants with higher gmax (also higher transpiration) require larger quantities of water to be moved across the rhizosphere, which results in a precipitous decrease in water potential at the soil-root interface, and therefore in the leaves. We demonstrated that the gmax -ψgs50 tradeoff can be predicted based on soil-plant hydraulics, and is impacted by plant hydraulic properties, such as plant hydraulic conductance, active root length and embolism resistance. We conclude that plants may therefore adjust their growth and/or their hydraulic properties to adapt to contrasting habitats and climate conditions.

Coordination of photosynthetic traits across soil and climate gradients.

"Least-cost theory" posits that C3 plants should balance rates of photosynthetic water loss and carboxylation in relation to the relative acquisition and maintenance costs of resources required for these activities. Here we investigated the dependency of photosynthetic traits on climate and soil properties using a new Australia-wide trait dataset spanning 528 species from 67 sites. We tested the hypotheses that plants on relatively cold or dry sites, or on relatively more fertile sites, would typically operate at greater CO2 drawdown (lower ratio of leaf internal to ambient CO2 , Ci :Ca ) during light-saturated photosynthesis, and at higher leaf N per area (Narea ) and higher carboxylation capacity (Vcmax 25 ) for a given rate of stomatal conductance to water vapour, gsw . These results would be indicative of plants having relatively higher water costs than nutrient costs. In general, our hypotheses were supported. Soil total phosphorus (P) concentration and (more weakly) soil pH exerted positive effects on the Narea -gsw and Vcmax 25 -gsw slopes, and negative effects on Ci :Ca . The P effect strengthened when the effect of climate was removed via partial regression. We observed similar trends with increasing soil cation exchange capacity and clay content, which affect soil nutrient availability, and found that soil properties explained similar amounts of variation in the focal traits as climate did. Although climate typically explained more trait variation than soil did, together they explained up to 52% of variation in the slope relationships and soil properties explained up to 30% of the variation in individual traits. Soils influenced photosynthetic traits as well as their coordination. In particular, the influence of soil P likely reflects the Australia's geologically ancient low-relief landscapes with highly leached soils. Least-cost theory provides a valuable framework for understanding trade-offs between resource costs and use in plants, including limiting soil nutrients.

Functional xylem characteristics associated with drought-induced embolism in angiosperms.

Hydraulic failure resulting from drought-induced embolism in the xylem of plants is a key determinant of reduced productivity and mortality. Methods to assess this vulnerability are difficult to achieve at scale, leading to alternative metrics and correlations with more easily measured traits. These efforts have led to the longstanding and pervasive assumed mechanistic link between vessel diameter and vulnerability in angiosperms. However, there are at least two problems with this assumption that requires critical re-evaluation: (1) our current understanding of drought-induced embolism does not provide a mechanistic explanation why increased vessel width should lead to greater vulnerability, and (2) the most recent advancements in nanoscale embolism processes suggest that vessel diameter is not a direct driver. Here, we review data from physiological and comparative wood anatomy studies, highlighting the potential anatomical and physicochemical drivers of embolism formation and spread. We then put forward key knowledge gaps, emphasising what is known, unknown and speculation. A meaningful evaluation of the diameter-vulnerability link will require a better mechanistic understanding of the biophysical processes at the nanoscale level that determine embolism formation and spread, which will in turn lead to more accurate predictions of how water transport in plants is affected by drought.

Age Dependent Partitioning Patterns of Essential Nutrients Induced by Copper Feeding Status in Leaves and Stems of Poplar.

Copper (Cu) is an essential micronutrient, and its deficiency can cause plants to undergo metabolic changes at several levels of organization. It has been shown that leaf age can play a role in nutrient partitioning along the shoot axis of poplar. In this study, we investigated the effect of Cu deficiency on the altered partitioning of essential macro and micronutrients in leaves and stems of different age. Cu deficiency was associated with higher concentrations of calcium, magnesium, sulfur, iron, zinc, manganese, and molybdenum in leaves and relatively higher concentrations of calcium, phosphorus, iron, and zinc in stems. Leaf and stem age had significant effects on nutrient partitioning. Principal component analyses revealed patterns that point to inverse influences in leaves and stems on nutrient partitioning. Specifically, these analyses revealed that nutrient partitioning in leaves was influenced by Cu feeding status more than developmental stage, whereas nutrient partitioning in stems was influenced by developmental stage more than Cu feeding status. These results suggest that Cu deficiency and developmental stage can significantly influence the partitioning and homeostasis of macro and micronutrients in poplar organs.

Physiological trait networks enhance understanding of crop growth and water use in contrasting environments.

Plant function arises from a complex network of structural and physiological traits. Explicit representation of these traits, as well as their connections with other biophysical processes, is required to advance our understanding of plant-soil-climate interactions. We used the Terrestrial Regional Ecosystem Exchange Simulator (TREES) to evaluate physiological trait networks in maize. Net primary productivity (NPP) and grain yield were simulated across five contrasting climate scenarios. Simulations achieving high NPP and grain yield in high precipitation environments featured trait networks conferring high water use strategies: deep roots, high stomatal conductance at low water potential ("risky" stomatal regulation), high xylem hydraulic conductivity and high maximal leaf area index. In contrast, high NPP and grain yield was achieved in dry environments with low late-season precipitation via water conserving trait networks: deep roots, high embolism resistance and low stomatal conductance at low leaf water potential ("conservative" stomatal regulation). We suggest that our approach, which allows for the simultaneous evaluation of physiological traits, soil characteristics and their interactions (i.e., networks), has potential to improve our understanding of crop performance in different environments. In contrast, evaluating single traits in isolation of other coordinated traits does not appear to be an effective strategy for predicting plant performance.

Recovery after deficiency: systemic copper prioritization and partitioning in the leaves and stems of hybrid poplar.

Copper (Cu) is important for many aspects of plant function including photosynthesis. It has been suggested that photosynthesis, especially in young leaves, is prioritized for Cu delivery after deficiency in hybrid poplar. To determine relative Cu delivery prioritization, we enriched hydroponic plant growth media of Cu-deficient poplar with 98% 65Cu and tracked Cu delivery after deficiency to young leaves, mature leaves and stems. Young leaves acquired ~58% more 65Cu on Day 1 and ~65% more 65Cu by Day 3 compared with mature leaves. Additionally, stomatal conductance (gs) was measured on leaves for 6 weeks and during a 3-day 65Cu pulse resupply period. During deficiency, mature leaves maintained a higher gs than younger leaves but 3 days after Cu resupply the younger leaves that had recovered showed the highest gs. In conclusion, these results provide a quantitative understanding of how Cu is systemically transported and distributed to photosynthetic and stem tissues.

Leaf nitrogen from the perspective of optimal plant function.

Leaf dry mass per unit area (LMA), carboxylation capacity (V cmax) and leaf nitrogen per unit area (Narea) and mass (Nmass) are key traits for plant functional ecology and ecosystem modelling. There is however no consensus about how these traits are regulated, or how they should be modelled. Here we confirm that observed leaf nitrogen across species and sites can be estimated well from observed LMA and V cmax at 25°C (V cmax25). We then test the hypothesis that global variations of both quantities depend on climate variables in specific ways that are predicted by leaf-level optimality theory, thus allowing both Narea to be predicted as functions of the growth environment.A new global compilation of field measurements was used to quantify the empirical relationships of leaf N to V cmax25 and LMA. Relationships of observed V cmax25 and LMA to climate variables were estimated, and compared to independent theoretical predictions of these relationships. Soil effects were assessed by analysing biases in the theoretical predictions.LMA was the most important predictor of Narea (increasing) and Nmass (decreasing). About 60% of global variation across species and sites in observed Narea, and 31% in Nmass, could be explained by observed LMA and V cmax25. These traits, in turn, were quantitatively related to climate variables, with significant partial relationships similar or indistinguishable from those predicted by optimality theory. Predicted trait values explained 21% of global variation in observed site-mean V cmax25, 43% in LMA and 31% in Narea. Predicted V cmax25 was biased low on clay-rich soils but predicted LMA was biased high, with compensating effects on Narea. Narea was overpredicted on organic soils. Synthesis. Global patterns of variation in observed site-mean Narea can be explained by climate-induced variations in optimal V cmax25 and LMA. Leaf nitrogen should accordingly be modelled as a consequence (not a cause) of V cmax25 and LMA, both being optimized to the environment. Nitrogen limitation of plant growth would then be modelled principally via whole-plant carbon allocation, rather than via leaf-level traits. Further research is required to better understand and model the terrestrial nitrogen and carbon cycles and their coupling.

Growth and grain yield of eight maize hybrids are aligned with water transport, stomatal conductance, and photosynthesis in a semi-arid irrigated system.

There is increasing interest in understanding how trait networks can be manipulated to improve the performance of crop species. Working towards this goal, we have identified key traits linking the acquisition of water, the transport of water to the sites of evaporation and photosynthesis, stomatal conductance, and growth across eight maize hybrid lines grown under well-watered and water-limiting conditions in Northern Colorado. Under well-watered conditions, hybrids with higher end-of-season growth and grain yield exhibited higher leaf-specific conductance, lower operating water potentials, higher rates of midday stomatal conductance, higher rates of net CO2 assimilation, and greater leaf osmotic adjustment. This trait network was similar under water-limited conditions with the notable exception that linkages between water transport, midday stomatal conductance, and growth were even stronger than under fully watered conditions. The maintenance of high leaf-specific conductance throughout the day was achieved via higher maximal conductance rates rather than lower susceptibility to conductance loss. Our results suggest that efforts to improve maize performance in well-watered and water-limiting conditions would benefit from considering the physiological trait networks governing water and carbon flux rather than focusing on single traits independently of one another.

Drought-Induced Root Pressure in Sorghum bicolor.

Root pressure, also manifested as profusive sap flowing from cut stems, is a phenomenon in some species that has perplexed biologists for much of the last century. It is associated with increased crop production under drought, but its function and regulation remain largely unknown. In this study, we investigated the initiation, mechanisms, and possible adaptive function of root pressure in six genotypes of Sorghum bicolor during a drought experiment in the greenhouse. We observed that root pressure was induced in plants exposed to drought followed by re-watering but possibly inhibited by 100% re-watering in some genotypes. We found that root pressure in drought stressed and re-watered plants was associated with greater ratio of fine: coarse root length and shoot biomass production, indicating a possible role of root allocation in creating root pressure and adaptive benefit of root pressure for shoot biomass production. Using RNA-Seq, we identified gene transcripts that were up- and down-regulated in plants with root pressure expression, focusing on genes for aquaporins, membrane transporters, and ATPases that could regulate inter- and intra-cellular transport of water and ions to generate positive xylem pressure in root tissue.

Weak tradeoff between xylem hydraulic efficiency and safety: climatic seasonality matters.

A classic theory proposes that plant xylem cannot be both highly efficient in water transport and resistant to embolism, and therefore a hydraulic efficiency-safety trade-off should exist. However, the trade-off is weak, and many species exhibit both low efficiency and low safety, falling outside of the expected trade-off space. It remains unclear under what climatic conditions these species could maintain competitive fitness. We compiled hydraulic efficiency and safety traits for 682 observations of 499 woody species from 178 sites world-wide and measured the position of each observation within the proposed trade-off space. For both angiosperms and gymnosperms, observations from sites with high climatic seasonality, especially precipitation seasonality, tended to have higher hydraulic safety and efficiency than observations from sites with low seasonality. Specifically, high vapour pressure deficit, high solar radiation, and low precipitation during the wet season were driving factors. Strong climatic seasonality and drought in both dry and wet seasons appear to be ecological filters that select for species with co-optimized safety and efficiency, whereas the opposite environmental conditions may allow the existence of plants with low efficiency and safety.

Increasing axial parenchyma fraction in the Malagasy Magnoliids facilitated the co-optimisation of hydraulic efficiency and safety.

The evolution of angiosperms was accompanied by the segregation and specialisation of their xylem tissues. This study aimed to determine whether the fraction and arrangement of parenchyma tissue influence the hydraulic efficiency-safety trade-off in the basal angiosperms. We examined xylem anatomical structure and hydraulic functioning of 28 woody species of Magnoliids in a tropical rainforest of Madagascar and reported, for the first time, quantitative measurements that support the relationship between vessel-to-xylem parenchyma connectivity and the hydraulic efficiency-safety trade-off. We also introduced a new measurement - the distance of species from the trade-off limit - to quantify the co-optimisation of hydraulic efficiency and safety. Although the basal angiosperms in this study had low hydraulic conductivity and safety, species with higher axial parenchyma fraction (APf) had significantly higher hydraulic conductivity. Hydraulic efficiency-safety optimisation was accompanied by higher APf and vessel-to-axial parenchyma connectivity. Conversely, species exhibiting high ray parenchyma fraction and high vessel-to-ray connectivity had lower Ks and were further away from the hydraulic trade-off limit line. Our results provide evidence that axial parenchyma fraction and paratracheal arrangement are associated with both enhanced hydraulic efficiency and safety.

Tip-to-base xylem conduit widening as an adaptation: causes, consequences, and empirical priorities.

In the stems of terrestrial vascular plants studied to date, the diameter of xylem water-conducting conduits D widens predictably with distance from the stem tip L approximating D âˆ Lb , with b ≈ 0.2. Because conduit diameter is central for conductance, it is essential to understand the cause of this remarkably pervasive pattern. We give reason to suspect that tip-to-base conduit widening is an adaptation, favored by natural selection because widening helps minimize the increase in hydraulic resistance that would otherwise occur as an individual stem grows longer and conductive path length increases. Evidence consistent with adaptation includes optimality models that predict the 0.2 exponent. The fact that this prediction can be made with a simple model of a single capillary, omitting much biological detail, itself makes numerous important predictions, e.g. that pit resistance must scale isometrically with conduit resistance. The idea that tip-to-base conduit widening has a nonadaptive cause, with temperature, drought, or turgor limiting the conduit diameters that plants are able to produce, is less consistent with the data than an adaptive explanation. We identify empirical priorities for testing the cause of tip-to-base conduit widening and underscore the need to study plant hydraulic systems leaf to root as integrated wholes.

What do you mean "functional" in ecology? Patterns versus processes.

Use of the term "functional" trait has increased exponentially in ecology. Although accounting for numerous ecological questions, this concept raises several issues. We propose that the term "functional" could be misleading because (1) no rigorous criteria exist to identify "functional" traits and (2) it suggests that only some traits ("functional" ones) can inform our understanding of species functioning, whatever the scale or discipline. Hence, the concept of "functional" trait in ecology is starting to be challenged and it remains unclear why some traits should be considered functional, whereas other traits should not. We argue that the most used "functional" traits are meaningful because they reflect important differences between populations or species, based on synchronic comparisons, that is, irrespective of time (hereafter "pattern" traits). Hence, they are useful for identifying trade-offs and strategies across large numbers of observations, usually at rather coarse scales, and are most often used in analyses of "big data." However, given that many ecological processes occur across short time scales and narrow gradients of climate and resource availability, the efficacy of these traits to inform us about these ecological processes appears questionable. We show that trait measurements that take time explicitly into account (hereafter "process" traits) differ from pattern traits because they quantify the flows of material and energy within a given environment across a defined period of time. Although pattern traits and process traits are both functional, it is important to understand the differences between the approaches. Moreover, better accounting of ontogeny, life form, plasticity, and genetic variability is required to enhance the convergence between pattern and process approaches. This revised framework allows more explicit connections between trait ecology and other biological sciences. It should enhance the study of processes at all scales in order to investigate efficiently the adaptive responses of biological organisms to climate change.

Wood day capacitance is related to water content, wood density, and anatomy across 30 temperate tree species.

Water released from wood during transpiration (capacitance) can meaningfully affect daily water use and drought response. To provide context for better understanding of capacitance mechanisms, we investigated links between capacitance and wood anatomy. On twigs of 30 temperate angiosperm tree species, we measured day capacitance (between predawn and midday), water content, wood density, and anatomical traits, that is, vessel dimensions, tissue fractions, and vessel-tissue contact fractions (fraction of vessel circumference in contact with other tissues). Across all species, wood density (WD) and predawn lumen volumetric water content (VWCL-pd ) together were the strongest predictors of day capacitance (r2adj = .44). Vessel-tissue contact fractions explained an additional ~10% of the variation in day capacitance. Regression models were not improved by including tissue lumen fractions. Among diffuse-porous species, VWCL-pd and vessel-ray contact fraction together were the best predictors of day capacitance, whereas among semi/ring-porous species, VWCL-pd , WD and vessel-fibre contact fraction were the best predictors. At predawn, wood was less than fully saturated for all species (lumen relative water content = 0.52 ± 0.17). Our findings imply that day capacitance depends on the amount of stored water, tissue connectivity and the bulk wood properties arising from WD (e.g., elasticity), rather than the fraction of any particular tissue.

Growing-season temperature and precipitation are independent drivers of global variation in xylem hydraulic conductivity.

Stem xylem-specific hydraulic conductivity (KS ) represents the potential for plant water transport normalized by xylem cross section, length, and driving force. Variation in KS has implications for plant transpiration and photosynthesis, growth and survival, and also the geographic distribution of species. Clarifying the global-scale patterns of KS and its major drivers is needed to achieve a better understanding of how plants adapt to different environmental conditions, particularly under climate change scenarios. Here, we compiled a xylem hydraulics dataset with 1,186 species-at-site combinations (975 woody species representing 146 families, from 199 sites worldwide), and investigated how KS varied with climatic variables, plant functional types, and biomes. Growing-season temperature and growing-season precipitation drove global variation in KS independently. Both the mean and the variation in KS were highest in the warm and wet tropical regions, and lower in cold and dry regions, such as tundra and desert biomes. Our results suggest that future warming and redistribution of seasonal precipitation may have a significant impact on species functional diversity, and is likely to be particularly important in regions becoming warmer or drier, such as high latitudes. This highlights an important role for KS in predicting shifts in community composition in the face of climate change.