Stem diameter growth rates in a fire‐prone savanna correlate with photosynthetic rate and branch‐scale biomass allocation,but not specific leaf area

Plant growth rates strongly determine ecosystem productivity and are a central element of plant ecological strategies. For laboratory and glasshouse‐grown seedlings, specific leaf area (SLA; ratio of leaf area to mass) is a key driver of interspecific variation in growth rate (GR). Consequently, SLA is often assumed to drive GR variation in field‐grown adult plants. However, there is an increasing evidence that this is not the general case. This suggests that GR – SLA relationships (and perhaps those for other traits) may vary depending on the age or size of the plants being studied. Here we investigated GR – trait relationships and their size dependence among 17 woody species from an open‐canopy, fire‐prone savanna in northern Australia. We tested the predictions that SLA and stem diameter growth rate would be positively correlated in saplings but unrelated in adults while, in both age classes, faster‐GR species would have higher light‐saturated photosynthetic rate (A_sat), higher leaf nutrient concentrations, higher branch‐scale biomass allocation to leaf versus stem tissues and lower wood density (WD). SLA showed no relationship to stem diameter GR, even in saplings, and the same was true of leaf N and P concentrations, and WD. However, branch‐scale leaf:stem allocation was strongly related to GR in both age groups, as was A_sat. Together, these two traits accounted for up to 80% of interspecific variation in adult GR, and 41% of sapling GR. A_sat is rarely measured in field‐based GR studies, and this is the first report of branch‐scale leaf:stem allocation (analogous to a benefit:cost ratio) in relation to plant growth rate. Our results suggest that we may yet find general trait‐drivers of field growth rates, but SLA will not be one.     

Relationships among ecologically important dimensions of plant trait variation in seven neotropical forests

BACKGROUND AND AIMS: When ecologically important plant traits are correlated they may be said to constitute an ecological 'strategy' dimension. Through identifying these dimensions and understanding their inter-relationships we gain insight into why particular trait combinations are favoured over others and into the implications of trait differences among species. Here we investigated relationships among several traits, and thus the strategy dimensions they represented, across 2134 woody species from seven Neotropical forests. METHODS: Six traits were studied: specific leaf area (SLA), the average size of leaves, seed and fruit, typical maximum plant height, and wood density (WD). Trait relationships were quantified across species at each individual forest as well as across the dataset as a whole. 'Phylogenetic' analyses were used to test for correlations among evolutionary trait-divergences and to ascertain whether interspecific relationships were biased by strong taxonomic patterning in the traits. KEY RESULTS: The interspecific and phylogenetic analyses yielded congruent results. Seed and fruit size were expected, and confirmed, to be tightly related. As expected, plant height was correlated with each of seed and fruit size, albeit weakly. Weak support was found for an expected positive relationship between leaf and fruit size. The prediction that SLA and WD would be negatively correlated was not supported. Otherwise the traits were predicted to be largely unrelated, being representatives of putatively independent strategy dimensions. This was indeed the case, although WD was consistently, negatively related to leaf size. CONCLUSIONS: The dimensions represented by SLA, seed/fruit size and leaf size were essentially independent and thus conveyed largely independent information about plant strategies. To a lesser extent the same was true for plant height and WD. Our tentative explanation for negative WD-leaf size relationships, now also known from other habitats, is that the traits are indirectly linked via plant hydraulics.     

Tree hydraulic traits are coordinated and strongly linked to climate‐of‐origin across a rainfall gradient

Plant hydraulic traits capture the impacts of drought stress on plant function, yet vegetation models lack sufficient information regarding trait coordination and variation with climate‐of‐origin across species. Here, we investigated key hydraulic and carbon economy traits of 12 woody species in Australia from a broad climatic gradient, with the aim of identifying the coordination among these traits and the role of climate in shaping cross‐species trait variation. The influence of environmental variation was minimized by a common garden approach, allowing us to factor out the influence of environment on phenotypic variation across species. We found that hydraulic traits (leaf turgor loss point, stomatal sensitivity to drought [P_gs], xylem vulnerability to cavitation [P_x], and branch capacitance [C_branch]) were highly coordinated across species and strongly related to rainfall and aridity in the species native distributional range. In addition, trade‐offs between drought tolerance and plant growth rate were observed across species. Collectively, these results provide critical insight into the coordination among hydraulic traits in modulating drought adaptation and will significantly advance our ability to predict drought vulnerability in these dominant trees species.     

Scale‐dependent trait correlations in a temperate tree community

Recent investigations of relationships among plant traits have generated important insights into plant form and function. However, relationships involving leaf area, leaf shape and plant height remain poorly resolved. Previous work has also focused on correlations between average trait values for individual species. It is unclear whether similar relationships occur within species. We searched for novel plant trait correlations by comparing leaf area, leaf circularity, specific leaf area (SLA) and plant height among 16 common woody plant species from a temperate forest in New Zealand. Analyses were conducted both within species (intra‐specifically) and among species (inter‐specifically) to determine whether trait correlations were scale dependent. Leaf area was unrelated to other leaf traits inter‐specifically. However, leaf area declined with plant height and increased with SLA intra‐specifically. Leaf circularity decreased with plant height inter‐specifically, but increased with plant height intra‐specifically. SLA increased with plant height both inter‐ and intra‐specifically. Leaf circularity increased with SLA inter‐specifically, but decreased with SLA intra‐specifically. Overall results showed that leaf shape, SLA and plant height are interrelated. However, intra‐specific relationships often differed substantially from inter‐specific relationships, suggesting that the processes shaping relationships between this suite of plant traits are scale‐dependent.     

Intraspecific trait variation can weaken interspecific trait correlations when assessing the whole‐plant economic spectrum

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.     

A trait‐based trade‐off between growth and mortality: evidence from 15 tropical tree species using size‐specific relative growth rates

A life‐history trade‐off between low mortality in the dark and rapid growth in the light is one of the most widely accepted mechanisms underlying plant ecological strategies in tropical forests. Differences in plant functional traits are thought to underlie these distinct ecological strategies; however, very few studies have shown relationships between functional traits and demographic rates within a functional group. We present 8 years of growth and mortality data from saplings of 15 species of Dipterocarpaceae planted into logged‐over forest in Malaysian Borneo, and the relationships between these demographic rates and four key functional traits: wood density, specific leaf area (SLA), seed mass, and leaf C:N ratio. Species‐specific differences in growth rates were separated from seedling size effects by fitting nonlinear mixed‐effects models, to repeated measurements taken on individuals at multiple time points. Mortality data were analyzed using binary logistic regressions in a mixed‐effects models framework. Growth increased and mortality decreased with increasing light availability. Species differed in both their growth and mortality rates, yet there was little evidence for a statistical interaction between species and light for either response. There was a positive relationship between growth rate and the predicted probability of mortality regardless of light environment, suggesting that this relationship may be driven by a general trade‐off between traits that maximize growth and traits that minimize mortality, rather than through differential species responses to light. Our results indicate that wood density is an important trait that indicates both the ability of species to grow and resistance to mortality, but no other trait was correlated with either growth or mortality. Therefore, the growth mortality trade‐off among species of dipterocarp appears to be general in being independent of species crossovers in performance in different light environments.     

Complex trait relationships between leaves and absorptive roots: Coordination in tissue N concentration but divergence in morphology

Leaves and absorptive roots (i.e., first‐order root) are above‐ and belowground plant organs related to resource acquisition; however, it is controversy over whether these two sets of functional traits vary in a coordinated manner. Here, we examined the relationships between analogous above‐ and belowground traits, including chemical (tissue C and N concentrations) and morphological traits (thickness and diameter, specific leaf area and root length, and tissue density) of 154 species sampling from eight subtropical and temperate forests. Our results showed that N concentrations of leaves and absorptive roots were positively correlated independent of phylogeny and plant growth forms, whereas morphological traits between above‐ and belowground organs varied independently. These results indicate that, different from plant economics spectrum theory, there is a complex integration of diverse adaptive strategies of plant species to above‐ and belowground environments, with convergent adaptation in nutrient traits but divergence in morphological traits across plant organs. Our results offer a new perspective for understanding the resource capture strategies of plants in adaptation to heterogeneous environments, and stress the importance of phylogenetic consideration in the discussion of cross‐species trait relationships.     

Within‐species patterns challenge our understanding of the leaf economics spectrum

The utility of plant functional traits for predictive ecology relies on our ability to interpret trait variation across multiple taxonomic and ecological scales. Using extensive data sets of trait variation within species, across species and across communities, we analysed whether and at what scales leaf economics spectrum (LES) traits show predicted trait–trait covariation. We found that most variation in LES traits is often, but not universally, at high taxonomic levels (between families or genera in a family). However, we found that trait covariation shows distinct taxonomic scale dependence, with some trait correlations showing opposite signs within vs. across species. LES traits responded independently to environmental gradients within species, with few shared environmental responses across traits or across scales. We conclude that, at small taxonomic scales, plasticity may obscure or reverse the broad evolutionary linkages between leaf traits, meaning that variation in LES traits cannot always be interpreted as differences in resource use strategy.     

Invasive plants in Minnesota are “joining the locals”: A trait‐based analysis

Questions

Predicting which newly arrived species will establish and become invasive is a problem that has long vexed researchers. In a study of cold temperate oak forest stands, we examined two contrasting hypotheses regarding plant functional traits to explain the success of certain non‐native species. Under the “join the locals” hypothesis, successful invaders are expected to share traits with resident species because they employ successful growth strategies under light‐limited understorey conditions. Instead, under the “try harder” hypothesis, successful invaders are expected to have traits different from native species in order to take advantage of unused niche space.

Location

Minnesota, USA.

Methods

We examined these two theories using 109 native and 11 non‐native plants in 68 oak forest stands. We focused on traits related to plant establishment and growth, including specific leaf area (SLA), leaf carbon‐to‐nitrogen ratio (C:N), wood density, plant maximum height, mycorrhizal type, seed mass and growth form. We compared traits of native and non‐native species using ordinations in multidimensional trait space and compared community‐weighted mean (CWM) trait values across sites.

Results

We found few differences between trait spaces occupied by native and non‐native species. Non‐native species occupied smaller areas of trait space than natives, yet were within that of the native species, indicating similar growth strategies. We observed a higher proportion of non‐native species in sites with higher native woody species CWM SLA and lower CWM C:N. Higher woody CWM SLA was observed in sites with higher soil pH, while lower CWM C:N was found in sites with higher light levels.

Conclusions

Non‐native plants in this system have functional traits similar to natives and are therefore “joining the locals.” However, non‐native plants may possess traits toward the acquisitive end of the native plant trait range, as evidenced by higher non‐native plant abundance in high‐resource environments.

     

Leaf hydraulic conductance varies with vein anatomy across Arabidopsis thaliana wild‐type and leaf vein mutants

Leaf venation is diverse across plant species and has practical applications from paleobotany to modern agriculture. However, the impact of vein traits on plant performance has not yet been tested in a model system such as Arabidopsis thaliana. Previous studies analysed cotyledons of A. thaliana vein mutants and identified visible differences in their vein systems from the wild type (WT). We measured leaf hydraulic conductance (K_leaf), vein traits, and xylem and mesophyll anatomy for A. thaliana WT (Col‐0) and four vein mutants (dot3‐111 and dot3‐134, and cvp1‐3 and cvp2‐1). Mutant true leaves did not possess the qualitative venation anomalies previously shown in the cotyledons, but varied quantitatively in vein traits and leaf anatomy across genotypes. The WT had significantly higher mean K_leaf. Across all genotypes, there was a strong correlation of K_leaf with traits related to hydraulic conductance across the bundle sheath, as influenced by the number and radial diameter of bundle sheath cells and vein length per area. These findings support the hypothesis that vein traits influence K_leaf, indicating the usefulness of this mutant system for testing theory that was primarily established comparatively across species, and supports a strong role for the bundle sheath in influencing K_leaf.     

Weak whole‐plant trait coordination in a seasonally dry South American stressful environment

A core question involving both plant physiology and community ecology is whether traits from different organs are coordinated across species, beyond pairwise trait correlations. The strength of within‐community trait coordination has been hypothesized to increase along gradients of environmental harshness, due to the cost of adopting ecological strategies out of the viable niche space supported by the abiotic conditions. We evaluated the strength of trait relationship and coordination in a stressful environment using 21 leaf and stem traits of 21 deciduous and evergreen woody species from a heath vegetation growing on coastal sandy plain in northeastern South America. The study region faces marked dry season, high soil salinity and acidity, and poor nutritional conditions. Results from multiple factor analyses supported two weak and independent axes of trait coordination, which accounted for 25%–29% of the trait variance using phylogenetically independent contrasts. Trait correlations on the multiple factor analyses main axis fit well with the global plant economic spectrum, with species investing in small leaves and dense stems as opposed to species with softer stems and large leaves. The species’ positions on the main functional axis corresponded to the competitor‐stress‐tolerant side of Grime's CSR triangle of plant strategies. The weak degree of trait coordination displayed by the heath vegetation species contradicted our expectation of high trait coordination in stressful environmental habitats. The distinct biogeographic origins of the species occurring in the study region and the prevalence of a regional environmental filter coupled with local homogeneous conditions could account for prevalence of trait independence we observed.     

Biogeographic constraints on the world‐wide leaf economics spectrum

Aim The world‐wide leaf economic spectrum (LES) describes tight coordination of leaf traits across global floras, reported to date as being largely independent of phylogeny and biogeography. Here, we present and test an alternative, historical perspective that predicts that biogeography places significant constraints on global trait evolution. These hypothesized constraints could lead to important deviations in leaf trait relationships between isolated floras that were influenced by different magnitudes of genetic constraint and selection. Location Global, including floristic regions of the Northern and Southern Hemispheres, eastern North America, East Asia (EAS), the Hawaiian Islands and tropical mainland floras. Methods We use a large leaf‐trait database (GLOPNET) and species native distribution data to test for variation in leaf trait relationships modulated by floristic region, controlling for climatic differences. Standardized major axis analyses were used to evaluate biogeographic effects on bivariate relationships between LES traits, including relationships of photosynthetic capacity and dark respiration rate (A_mass–R_d‐mass), leaf lifespan and mass per area ratio (LL–LMA), and photosynthetic capacity and nitrogen content (A_mass–N_mass). Results Independent of climate or biome, floras of different evolutionary histories exhibited different leaf trait allometries. Floras of the Northern Hemisphere exhibited greater rates of return on resource investment (steeper slopes for the trait relationships analysed), and the more diverse temperate EAS flora exhibited greater slopes or intercepts in leaf trait relationships, with the exception of the A_mass–N_mass relationship. In contrast to our hypothesis, plants of the floristically isolated Hawaiian Islands exhibited a similar A_mass–N_mass relationship to those of mainland tropical regions. Main conclusions Differences in leaf trait allometries among global floristic regions support a historical perspective in understanding leaf trait relationships and suggest that independent floras can exhibit different tradeoffs in resource capture strategies.     

Plasticity in above‐ and belowground resource acquisition traits in response to single and multiple environmental factors in three tree species

Functional trait plasticity is a major component of plant adjustment to environmental stresses. Here, we explore how multiple local environmental gradients in resources required by plants (light, water, and nutrients) and soil disturbance together influence the direction and amplitude of intraspecific changes in leaf and fine root traits that facilitate capture of these resources. We measured population‐level analogous above‐ and belowground traits related to resource acquisition, i.e. “specific leaf area”–“specific root length” (SLA–SRL), and leaf and root N, P, and dry matter content (DMC), on three dominant understory tree species with contrasting carbon and nutrient economics across 15 plots in a temperate forest influenced by burrowing seabirds. We observed similar responses of the three species to the same single environmental influences, but partially species‐specific responses to combinations of influences. The strength of intraspecific above‐ and belowground trait responses appeared unrelated to species resource acquisition strategy. Finally, most analogous leaf and root traits (SLA vs. SRL, and leaf versus root P and DMC) were controlled by contrasting environmental influences. The decoupled responses of above‐ and belowground traits to these multiple environmental factors together with partially species‐specific adjustments suggest complex responses of plant communities to environmental changes, and potentially contrasting feedbacks of plant traits with ecosystem properties. We demonstrate that despite the growing evidence for broadly consistent resource‐acquisition strategies at the whole plant level among species, plants also show partially decoupled, finely tuned strategies between above‐ and belowground parts at the intraspecific level in response to their environment. This decoupling within species suggests a need for many species‐centred ecological theories on how plants respond to their environments (e.g. competitive/stress‐tolerant/ruderal and response‐effect trait frameworks) to be adapted to account for distinct plant‐environment interactions among distinct individuals of the same species and parts of the same individual.     

Phylogenetic structural equation modelling reveals no need for an ‘origin’ of the leaf economics spectrum

The leaf economics spectrum (LES) is a prominent ecophysiological paradigm that describes global variation in leaf physiology across plant ecological strategies using a handful of key traits. Nearly a decade ago, Shipley et al. (2006) used structural equation modelling to explore the causal functional relationships among LES traits that give rise to their strong global covariation. They concluded that an unmeasured trait drives LES covariation, sparking efforts to identify the latent physiological trait underlying the ‘origin’ of the LES. Here, we use newly developed phylogenetic structural equation modelling approaches to reassess these conclusions using both global LES data as well as data collected across scales in the genus Helianthus. For global LES data, accounting for phylogenetic non‐independence indicates that no additional unmeasured traits are required to explain LES covariation. Across datasets in Helianthus, trait relationships are highly variable, indicating that global‐scale models may poorly describe LES covariation at non‐global scales.     

A global meta‐analysis of the relative extent of intraspecific trait variation in plant communities

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta‐analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole‐plant (e.g. plant height) vs. organ‐level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait‐based community and ecosystem studies.     

Trade-offs between plant leaf hydraulic and economic traits

Leaf is the most important organ for carbon-water coupling of a plant because it is the primary medium for photosynthesis. It also acts as the hydraulic bottleneck and safety valve against hydraulic catastrophic dysfunctions. The leaf economics spectrum, which reflects the balance between investments and returns of leaf economic traits, provides a useful framework for examining species strategies as shaped by their evolutionary history. Changes in leaf hydraulic traits will influence leaf economic traits as well as plant survival and growth. Exploring trade-offs between leaf hydraulic and economic traits is thus of significance for modeling carbon-water relations, understanding the mechanisms of water/carbon investments, and extending the leaf economic spectrum. In this review, we first examined the trade-offs between leaf hydraulic and economic traits. Specially, we analyzed the relationships between leaf hydraulic conductivity and hydraulic vulnerability, water potential at the turgor loss point, water capacitance, safety margin, and leaf morphological, structural and functional traits. We then discussed potential mechanisms regulating leaf hydraulic and economic traits from leaf morphology, anatomy, venation, and stomatal functions. Finally, we proposed future research to: (1) develop an integrated whole-plant economics spectrum, including carbon-nitrogen-water resources and root-stem-leaf hydraulic transport system that will help revealing ecophysiological mechanisms of plant structure-functional coupling, carbon sequestration and water use; (2) explore a generalized trade-offs among leaf hydraulic safety, hydraulic efficiency and carbon fixation efficiency to advance our understanding of the relationships between biophysical structure and physiological metabolism in plant leaf construction under drought stress; and (3) explore the carbon-water metabolic relationship and coupling of water transport and growth rate for the metabolic theory and predictions at community scale.     

Correlated evolution of stem and leaf hydraulic traits in Pereskia (Cactaceae)

Recent studies have demonstrated significant correlations between stem and leaf hydraulic properties when comparing across species within ecological communities. This implies that these traits are co-evolving, but there have been few studies addressing plant water relations within an explicitly evolutionary framework. This study tests for correlated evolution among a suite of plant water-use traits and environmental parameters in seven species of Pereskia (Cactaceae), using phylogenetically independent contrasts. There were significant evolutionary correlations between leaf-specific xylem hydraulic conductivity, Huber Value, leaf stomatal pore index, leaf venation density and leaf size, but none of these traits appeared to be correlated with environmental water availability; only two water relations traits - mid-day leaf water potentials and photosynthetic water use efficiency - correlated with estimates of moisture regime. In Pereskia, it appears that many stem and leaf hydraulic properties thought to be critical to whole-plant water use have not evolved in response to habitat shifts in water availability. This may be because of the extremely conservative stomatal behavior and particular rooting strategy demonstrated by all Pereskia species investigated. These results highlight the need for a lineage-based approach to understand the relative roles of functional traits in ecological adaptation.     

植物叶片水力与经济性状权衡关系的研究进展

叶片既是植物光合产物形成的主要场所, 又是整株植物的水力瓶颈、应对灾难性水力失调的安全阀门, 是植物碳水耦合权衡的重要器官。叶经济型谱反映了叶片经济性状“投资-收益”的权衡, 为验证植物进化过程中形成的物种对策提供了适用的理论框架。叶片水力性状变化会影响叶片经济性状及植物存活和生长。因此, 探索植物叶片水力与经济性状的权衡关系, 对建立植物碳-水耦合模型、揭示植物水-碳投资机理、扩展植物性状型谱等均有重要意义。该文首先综述了叶片水力性状、经济性状及两者之间的权衡关系, 分析了叶片导水率与水力脆弱性、失膨点水势、水容、安全阈值等水力性状以及与叶片的形态、结构和气体交换功能性状之间的关系。然后, 从叶片形态、解剖和叶脉网络结构以及气孔功能方面探讨了叶片水力性状与经济性状的调节机制。最后, 提出今后应加强三方面的研究: (1)探索建立植物根-茎-叶水力输导系统的碳-氮-水资源的整株经济型谱, 以揭示植物功能结构耦合、高效固碳用水的生理生态学机制; (2)探索叶片水力安全、水力效率和固碳效率之间的普适性权衡关系, 以深入理解抗旱植物叶片构建的生物物理结构与生理代谢的关系; (3)探索个体水平碳水代谢关系、水分运输与生长速率的耦合, 为代谢推演理论和植物群落尺度预测提供基础。     

Bottom‐up and top‐down effects of tree species diversity on leaf insect herbivory

The diversity of plant neighbors commonly results in direct, bottom‐up effects on herbivore ability to locate their host, and in indirect effects on herbivores involving changes in plant traits and a top‐down control by their enemies. Yet, the relative contribution of bottom‐up and top‐down forces remains poorly understood. We also lack knowledge on the effect of abiotic constraints such as summer drought on the strength and direction of these effects. We measured leaf damage on pedunculate oak (Quercus robur), alone or associated with birch, pine or both in a long‐term tree diversity experiment (ORPHEE), where half of the plots were irrigated while the other half remained without irrigation and received only rainfall. We tested three mechanisms likely to explain the effects of oak neighbors on herbivory: (1) Direct bottom‐up effects of heterospecific neighbors on oak accessibility to herbivores, (2) indirect bottom‐up effects of neighbors on the expression of leaf traits, and (3) top‐down control of herbivores by predators. Insect herbivory increased during the growth season but was independent of neighbor identity and irrigation. Specific leaf area, leaf toughness, and thickness varied with neighbor identity while leaf dry matter content or C:N ratio did not. When summarized in a principal component analysis (PCA), neighbor identity explained 87% of variability in leaf traits. PCA axes partially predicted herbivory. Despite greater rates of attack on dummy caterpillars in irrigated plots, avian predation, and insect herbivory remained unrelated. Our study suggests that neighbor identity can indirectly influence insect herbivory in mixed forests by modifying leaf traits. However, we found only partial evidence for these trait‐mediated effects and suggest that more attention should be paid to some unmeasured plant traits such as secondary metabolites, including volatile organic compounds, to better anticipate the effects of climate change on plant‐insect interactions in the future.     

Cross-species patterns in the coordination between leaf and stem traits, and their implications for plant hydraulics

Through identifying and understanding ecologically important dimensions of plant trait variation we gain insight into why particular trait combinations are favoured and into the implications of trait differences among species. Here, we describe relationships among several poorly understood leaf and stem traits across species from several Australian vegetation types. Species with lower wood density (WD) consistently deployed more leaf area per unit shoot mass (LA/SM), as did the larger-leaved species within forested sites. Higher LA/SM is likely to lead to faster growth rates, implying a previously unrecognized implication to interspecific variation in leaf size and WD. Leaf : sapwood area ratio is one of several important traits contributing to a plant's water-use strategy, yet, we still only poorly understand how plants vary in the extent to which hydraulic properties and traits such as leaf size, WD and LM/SM are coordinated, and what the implications of this variation may be.