Characterizing tree trait variance over spatiotemporal scales.

Beyond the study of the mean, functional ecology lacks a concise characterization of trait variance patterns across spatiotemporal scales. Traits are measured in different ways, using different metrics, and at different spatial (and rarely temporal) scales. This study expands on previous research by applying a ubiquitous and widely used empirical model-Taylor's Power Law-to functional trait variance with the goal of identifying general patterns of trait variance scaling (the behavior of trait variance across scales). We compiled data on tree seedling communities monitored over 10 years across 213 2 m2 plots and functional trait data from a subtropical forest in Puerto Rico. We examined trait-based Taylor's Power Law at nested spatial and temporal scales. The scaling of variance with the mean was idiosyncratic across traits suggesting that the drivers of variation are likely to differ across traits that may make variance scaling theory elusive. However, slopes varied more in space than through time, suggesting that spatial environmental variability may have a larger role in driving trait variance than temporal variability. Empirical models that characterize taxonomic patterns across spatiotemporal scales, like Taylor's Power Law, can provide an insight into the scaling of functional traits, a necessary next step toward a more predictive trait-based ecology.

Understanding Drivers of Variation and Predicting Variability Across Levels of Biological Organization.

Differences within a biological system are ubiquitous, creating variation in nature. Variation underlies all evolutionary processes and allows persistence and resilience in changing environments; thus, uncovering the drivers of variation is critical. The growing recognition that variation is central to biology presents a timely opportunity for determining unifying principles that drive variation across biological levels of organization. Currently, most studies that consider variation are focused at a single biological level and not integrated into a broader perspective. Here we explain what variation is and how it can be measured. We then discuss the importance of variation in natural systems, and briefly describe the biological research that has focused on variation. We outline some of the barriers and solutions to studying variation and its drivers in biological systems. Finally, we detail the challenges and opportunities that may arise when studying the drivers of variation due to the multi-level nature of biological systems. Examining the drivers of variation will lead to a reintegration of biology. It will further forge interdisciplinary collaborations and open opportunities for training diverse quantitative biologists. We anticipate that these insights will inspire new questions and new analytic tools to study the fundamental questions of what drives variation in biological systems and how variation has shaped life.

Beyond leaf habit: generalities in plant function across 97 tropical dry forest tree species.

Leaf habit has been hypothesized to define a linkage between the slow-fast plant economic spectrum and the drought resistance-avoidance trade-off in tropical forests ('slow-safe vs fast-risky'). However, variation in hydraulic traits as a function of leaf habit has rarely been explored for a large number of species. We sampled leaf and branch functional traits of 97 tropical dry forest tree species from four sites to investigate whether patterns of trait variation varied consistently in relation to leaf habit along the 'slow-safe vs fast-risky' trade-off. Leaf habit explained from 0% to 43.69% of individual trait variation. We found that evergreen and semi-deciduous species differed in their location along the multivariate trait ordination when compared to deciduous species. While deciduous species showed consistent trait values, evergreen species trait values varied as a function of the site. Last, trait values varied in relation to the proportion of deciduous species in the plant community. We found that leaf habit describes the strategies that define drought avoidance and plant economics in tropical trees. However, leaf habit alone does not explain patterns of trait variation, which suggests quantifying site-specific or species-specific uncertainty in trait variation as the way forward.

Trait-based signatures of cloud base height in a tropical cloud forest.

PREMISE: Clouds have profound consequences for ecosystem structure and function. Yet, the direct monitoring of clouds and their effects on biota is challenging especially in remote and topographically complex tropical cloud forests. We argue that known relationships between climate and the taxonomic and functional composition of plant communities may provide a fingerprint of cloud base height, thus providing a rapid and cost-effective assessment in remote tropical cloud forests. METHODS: To detect cloud base height, we compared species turnover and functional trait values among herbaceous and woody plant communities in an ecosystem dominated by cloud formation. We measured soil and air temperature, soil nutrient concentrations, and extracellular enzyme activity. We hypothesized that woody and herbaceous plants would provide signatures of cloud base height, as evidenced by abrupt shifts in both taxonomic composition and plant function. RESULTS: We demonstrated abrupt changes in taxonomic composition and the community- weighted mean of a key functional trait, specific leaf area, across elevation for both woody and herbaceous species, consistent with our predictions. However, abrupt taxonomic and functional changes occurred 100 m higher in elevation for herbaceous plants compared to woody ones. Soil temperature abruptly decreased where herbaceous taxonomic and functional turnover was high. Other environmental variables including soil biogeochemistry did not explain the abrupt change observed for woody plant communities. CONCLUSIONS: We provide evidence that a trait-based approach can be used to estimate cloud base height. We outline how rises in cloud base height and differential environmental requirements between growth forms can be distinguished using this approach.

Climate shapes and shifts functional biodiversity in forests worldwide.

Much ecological research aims to explain how climate impacts biodiversity and ecosystem-level processes through functional traits that link environment with individual performance. However, the specific climatic drivers of functional diversity across space and time remain unclear due largely to limitations in the availability of paired trait and climate data. We compile and analyze a global forest dataset using a method based on abundance-weighted trait moments to assess how climate influences the shapes of whole-community trait distributions. Our approach combines abundance-weighted metrics with diverse climate factors to produce a comprehensive catalog of trait-climate relationships that differ dramatically-27% of significant results change in sign and 71% disagree on sign, significance, or both-from traditional species-weighted methods. We find that (i) functional diversity generally declines with increasing latitude and elevation, (ii) temperature variability and vapor pressure are the strongest drivers of geographic shifts in functional composition and ecological strategies, and (iii) functional composition may currently be shifting over time due to rapid climate warming. Our analysis demonstrates that climate strongly governs functional diversity and provides essential information needed to predict how biodiversity and ecosystem function will respond to climate change.

Contrasting patterns of leaf trait variation among and within species during tropical dry forest succession in Costa Rica.

A coordinated response to environmental drivers amongst individual functional traits is central to the plant strategy concept. However, whether the trait co-ordination observed at the global scale occurs at other ecological scales (especially within species) remains an open question. Here, for sapling communities of two tropical dry forest types in Costa Rica, we show large differences amongst traits in the relative contribution of species turnover and intraspecific variation to their directional changes in response to environmental changes along a successional gradient. We studied the response of functional traits associated with the leaf economics spectrum and drought tolerance using intensive sampling to analyse inter- and intra-specific responses to environmental changes and ontogeny. Although the overall functional composition of the sapling communities changed during succession more through species turnover than through intraspecific trait variation, their relative contributions differed greatly amongst traits. For instance, community mean specific leaf area changed mostly due to intraspecific variation. Traits of the leaf economics spectrum showed decoupled responses to environmental drivers and ontogeny. These findings emphasise how divergent ecological mechanisms combine to cause great differences in changes of individual functional traits over environmental gradients and ecological scales.

Organismal responses to habitat change: herbivore performance, climate and leaf traits in regenerating tropical dry forests.

The ecological effects of large-scale climate change have received much attention, but the effects of the more acute form of climate change that results from local habitat alteration have been less explored. When forest is fragmented, cut, thinned, cleared or otherwise altered in structure, local climates and microclimates change. Such changes can affect herbivores both directly (e.g. through changes in body temperature) and indirectly (e.g. through changes in host plant traits). We advance an eco-physiological framework to understand the effects of changing forests on herbivorous insects. We hypothesize that if tropical forest caterpillars are climate and resource specialists, then they should have reduced performance outside of mature forest conditions. We tested this hypothesis with a field experiment contrasting the performance of Rothschildia lebeau (Saturniidae) caterpillars feeding on the host plant Casearia nitida (Salicaceae) in two different aged and structured tropical dry forests in Area de Conservación Guanacaste, Costa Rica. Compared to more mature closed-canopy forest, in younger secondary forest we found that: (1) ambient conditions were hotter, drier and more variable; (2) caterpillar growth and development were reduced; and (3) leaves were tougher, thicker and drier. Furthermore, caterpillar growth and survival were negatively correlated with these leaf traits, suggesting indirect host-mediated effects of climate on herbivores. Based on the available evidence, and relative to mature forest, we conclude that reduced herbivore performance in young secondary forest could have been driven by changes in climate, leaf traits (which were likely climate induced) or both. However, additional studies will be needed to provide more direct evidence of cause-and-effect and to disentangle the relative influence of these factors on herbivore performance in this system.

Tree height-diameter allometry across the United States.

The relationship between tree height and diameter is fundamental in determining community and ecosystem structure as well as estimates of biomass and carbon storage. Yet our understanding of how tree allometry relates to climate and whole organismal function is limited. We used the Forest Inventory and Analysis National Program database to determine height-diameter allometries of 2,976,937 individuals of 293 tree species across the United States. The shape of the allometric relationship was determined by comparing linear and nonlinear functional forms. Mixed-effects models were used to test for allometric differences due to climate and floristic (between angiosperms and gymnosperms) and functional groups (leaf habit and shade tolerance). Tree allometry significantly differed across the United States largely because of climate. Temperature, and to some extent precipitation, in part explained tree allometric variation. The magnitude of allometric variation due to climate, however, had a phylogenetic signal. Specifically, angiosperm allometry was more sensitive to differences in temperature compared to gymnosperms. Most notably, angiosperm height was more negatively influenced by increasing temperature variability, whereas gymnosperm height was negatively influenced by decreasing precipitation and increasing altitude. There was little evidence to suggest that shade tolerance influenced tree allometry except for very shade-intolerant trees which were taller for any given diameter. Tree allometry is plastic rather than fixed and scaling parameters vary around predicted central tendencies. This allometric variation provides insight into life-history strategies, phylogenetic history, and environmental limitations at biogeographical scales.

Interannual variability of growth and reproduction in Bursera simaruba: the role of allometry and resource variability.

Plants are expected to differentially allocate resources to reproduction, growth, and survival in order to maximize overall fitness. Life history theory predicts that the allocation of resources to reproduction should occur at the expense of vegetative growth. Although it is known that both organism size and resource availability can influence life history traits, few studies have addressed how size dependencies of growth and reproduction and variation in resource supply jointly affect the coupling between growth and reproduction. In order to understand the relationship between growth and reproduction in the context of resource variability, we utilize a long-term observational data set consisting of 670 individual trees over a 10-year period within a local population of Bursera simaruba (L.) Sarg. We (1) quantify the functional form and variability in the growth-reproduction relationship at the population and individual-tree level and (2) develop a theoretical framework to understand the allometric dependence of growth and reproduction. Our findings suggest that the differential responses of allometric growth and reproduction to resource availability, both between years and between microsites, underlie the apparent relationship between growth and reproduction. Finally, we offer an alternative approach for quantifying the relationship between growth and reproduction that accounts for variation in allometries.

The return of the variance: intraspecific variability in community ecology.

Despite being recognized as a promoter of diversity and a condition for local coexistence decades ago, the importance of intraspecific variance has been neglected over time in community ecology. Recently, there has been a new emphasis on intraspecific variability. Indeed, recent developments in trait-based community ecology have underlined the need to integrate variation at both the intraspecific as well as interspecific level. We introduce new T-statistics ('T' for trait), based on the comparison of intraspecific and interspecific variances of functional traits across organizational levels, to operationally incorporate intraspecific variability into community ecology theory. We show that a focus on the distribution of traits at local and regional scales combined with original analytical tools can provide unique insights into the primary forces structuring communities.