Performance-based inference of selection on stomatal length and specific leaf area varies with climate-of-origin of the forest tree, Eucalyptus ovata

Understanding how functional traits affect plant performance and fitness is a key step in unravelling the role of natural selection in shaping the evolutionary trajectory of populations. We examined early-age selection acting on leaf traits via their effects on growth performance and fitness, measured in Eucalyptus ovata trees planted in a common-garden field trial embedded in a reforestation planting in Tasmania, Australia. We focused on two important leaf traits - stomatal length and specific leaf area (SLA) - measured two years after planting, and compared interplanted E. ovata groups originating from dry and wet home-site climates, with the trial site having intermediate long-term mean annual rainfall. Two-year height growth was used as the performance attribute, and the time-averaged tree survival over the subsequent six years as the fitness component. There was evidence for performance-based selection on the leaf traits, with the strength and form of selection depending on the trait and climate group being considered. In this sense, selection in the dry group operated mainly on stomatal length where a combination of directional (favouring longer stomata) and stabilizing selection was detected, whereas selection in the wet group acted only on SLA and was purely stabilizing. Estimates of performance-based correlational selection were not statistically significant. For both climate groups, estimates of fitness-based selection gradients provided evidence for significant directional (but not quadratic) selection on height performance, favouring individuals with faster growth, but did not indicate statistical support for direct effects of the leaf traits on tree survival, conditional on measured performance. These results validated qualitative inferences of selection from the performance-based analysis, and suggested that selection on the leaf traits appeared to be mediated by their effects on early-age height performance, which in turn directly influenced later-age survival. We discuss the mechanisms by which the focal traits may have affected height performance, and likely factors contributing to the different patterns of phenotypic selection observed in the two groups experiencing the same environment. We also provide expressions of analytical derivatives that were developed for the estimation of selection gradients based on a logistic regression model relating a binary fitness response to linear and nonlinear covariate terms for the target regressor variables.info:eu-repo/semantics/publishedVersio

Population divergence along a genetic line of least resistance in the tree species Eucalyptus globulus

The evolutionary response to selection depends on the distribution of genetic variation in traits under selection within populations, as defined by the additive genetic variance-covariance matrix (G). The structure and evolutionary stability of G will thus influence the course of phenotypic evolution. However, there are few studies assessing the stability of G and its relationship with population divergence within foundation tree species. We compared the G-matrices of Mainland and Island population groups of the forest tree Eucalyptus globulus, and determined the extent to which population divergence aligned with within-population genetic (co)variation. Four key wood property traits exhibiting signals of divergent selection were studied—wood density, extractive content, and lignin content and composition. The comparison of G-matrices of the mainland and island populations indicated that the G-eigenstructure was relatively well preserved at an intra-specific level. Population divergence tended to occur along a major direction of genetic variation in G. The observed conservatism of G, the moderate evolutionary timescale, and close relationship between genetic architecture and population trajectories suggest that genetic constraints may have influenced the evolution and diversification of the E. globulus populations for the traits studied. However, alternative scenarios, including selection aligning genetic architecture and population divergence, are discussedinfo:eu-repo/semantics/publishedVersio

Linking leaf economic and hydraulic traits with early-age growth performance and survival of Eucalyptus pauciflora

Selection on plant functional traits may occur through their direct effects on fitness (or a fitness component), or may be mediated by attributes of plant performance which have a direct impact on fitness. Understanding this link is particularly challenging for long-lived organisms, such as forest trees, where lifetime fitness assessments are rarely achievable, and performance features and fitness components are usually quantified from early-life history stages. Accordingly, we studied a cohort of trees from multiple populations of Eucalyptus pauciflora grown in a common-garden field trial established at the hot and dry end of the species distribution on the island of Tasmania, Australia. We related the within-population variation in leaf economic (leaf thickness, leaf area and leaf density) and hydraulic (stomatal density, stomatal length and vein density) traits, measured from two-year-old plants, to two-year growth performance (height and stem diameter) and to a fitness component (seven-year survival). When performance-trait relationships were modelled for all traits simultaneously, statistical support for direct effects on growth performance was only observed for leaf thickness and leaf density. Performance-based estimators of directional selection indicated that individuals with reduced leaf thickness and increased leaf density were favoured. Survival-performance relationships were consistent with size- dependent mortality, with fitness-based selection gradients estimated for performance measures providing evidence for directional selection favouring individuals with faster growth. There was no statistical support for an effect associated with the fitness-based quadratic selection gradient estimated for growth performance. Conditional on a performance measure, fitness-based directional selection gradients estimated for the leaf traits did not provide statistical support for direct effects of the focal traits on tree survival. This suggested that, under the environmental conditions of the trial site and time period covered in the current study, early-stage selection on the studied leaf traits may be mediated by their effects on growth performance, which in turn has a positive direct influence on later-age survival. We discuss the potential mechanistic basis of the direct effects of the focal leaf traits on tree growth, and the relevance of a putative causal pathway of trait effects on fitness through mediation by growth performance in the studied hot and dry environmentinfo:eu-repo/semantics/publishedVersio

Shifts in Species Interactions Due to the Evolution of Functional Differences between Endemics and Non-Endemics: An Endemic Syndrome Hypothesis

Species ranges have been shifting since the Pleistocene, whereby fragmentation, isolation, and the subsequent reduction in gene flow have resulted in local adaptation of novel genotypes and the repeated evolution of endemic species. While there is a wide body of literature focused on understanding endemic species, very few studies empirically test whether or not the evolution of endemics results in unique function or ecological differences relative to their widespread congeners; in particular while controlling for environmental variation. Using a common garden composed of 15 Eucalyptus species within the subgenus Symphyomyrtus (9 endemic to Tasmania, 6 non-endemic), here we hypothesize and show that endemic species are functionally and ecologically different from non-endemics. Compared to non-endemics, endemic Eucalyptus species have a unique suite of functional plant traits that have extended effects on herbivores. We found that while endemics occupy many diverse habitats, they share similar functional traits potentially resulting in an endemic syndrome of traits. This study provides one of the first empirical datasets analyzing the functional differences between endemics and non-endemics in a common garden setting, and establishes a foundation for additional studies of endemic/non-endemic dynamics that will be essential for understanding global biodiversity in the midst of rapid species extinctions and range shifts as a consequence of global change

Directional Selection on Tree Seedling Traits Driven by Experimental Drought Differs Between Mesic and Dry Populations

Original ResearchWe evaluated population differences and drought-induced phenotypic selection on four seedling traits of the Australian forest tree Eucalyptus pauciflora using a glasshouse drydown experiment. We compared dry and mesic populations and tested for directional selection on lamina length (reflecting leaf size), leaf shape, the node of ontogenetic transition to the petiolate leaf (reflecting the loss of vegetative juvenility), and lignotuber size (reflecting a recovery trait). On average, the dry population had smaller and broader leaves, greater retention of the juvenile leaf state and larger lignotubers than the mesic population, but the populations did not differ in seedling survival. While there was statistical support for directional selection acting on the focal traits in one or other population, and for differences between populations in selection gradient estimates for two traits, only one trait—lamina length—exhibited a pattern of directional selection consistent with the observed population differences being a result of past adaptation to reduce seedling susceptibility to acute drought. The observed directional selection for lamina length in the mesic population suggests that future increases in drought risk in the wild will shift the mean of the mesic population toward that of the dry population. Further, we provide evidence suggesting an early age trade-off between drought damage and recovery traits, with phenotypes which develop larger lignotubers early being more susceptible to drought death. Such trade-offs could have contributed to the absence of population mean differences in survival, despite marked differentiation in seedling traitsinfo:eu-repo/semantics/publishedVersio

Genetic control of the operculum and capsule morphology of Eucalyptus globulus

The petaline operculum that covers the inner whorls until anthesis and the woody capsule that develops after fertilization are reproductive structures of eucalypts that protect the flower and seeds. Although they are distinct organs, they both develop from flower buds and this common ontogeny suggests shared genetic control. In Eucalyptus globulus their morphology is variable and we aimed to identify the quantitative trait loci (QTL) underlying this variation and determine whether there is common genetic control of these ecologically and taxonomically important reproductive structures.EEA Bella VistaFil: Hernández, Mariano Agustín. University of Tasmania. School of Natural Sciences; Australia.Fil: Hernández, Mariano Agustín. University of Tasmania. ARC Training Centre for Forest Value; Australia.Fil: Hernández, Mariano Agustín. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Bella Vista; Argentina.Fil: Butler, Jakob B. University of Tasmania. School of Natural Sciences; Australia.Fil: Ammitzboll, Hans. University of Tasmania. School of Natural Sciences; Australia.Fil: Ammitzboll, Hans. University of Tasmania. ARC Training Centre for Forest Value; Australia.Fil: Weller, James L. University of Tasmania. School of Natural Sciences; Australia.Fil: Weller, James L. Australian Research Council Centre of Excellence for Plant Success in Nature and Agriculture; Australia.Fil: Vaillancourt, René E. University of Tasmania. School of Natural Sciences; Australia.Fil: Vaillancourt, René E. University of Tasmania. ARC Training Centre for Forest Value; Australia.Fil: Potts, Brad M. University of Tasmania. School of Natural Sciences; Australia.Fil: Potts, Brad M. University of Tasmania. ARC Training Centre for Forest Value; Australia

QTL analysis for growth and wood properties across multiple pedigrees and sites in Eucalyptus globulus

Eucalyptus globulus is the most widely planted species for pulpwood production in temperate regions of the world and there are breeding programs in numerous countries. There is interest in molecular approaches to breeding, particularly marker assisted selection of wood properties. QTL analysis has an important role in identifying positional candidate genes responsible for variation in wood properties. This is one approach to targeting genes which may harbour functional allelic variants (SNPs). The objective of this study was to detect and validate QTL across multiple sites and pedigrees, in order to identify genomic regions and genes affecting growth and wood properties with wide applicability in the species. We also aimed to determine the proportion of QTL which were stable in their expression across sites of contrasting productivity. Such information will be important to exploit the full potential of the impending Eucalyptus genome sequences. [Oral Presentation

Assessing a Bayesian Approach for Detecting Exotic Hybrids between Plantation and Native Eucalypts

Eucalyptus globulus is grown extensively in plantations outside its native range in Australia. Concerns have been raised that the species may pose a genetic risk to native eucalypt species through hybridisation and introgression. Methods for identifying hybrids are needed to enable assessment and management of this genetic risk. This paper assesses the efficiency of a Bayesian approach for identifying hybrids between the plantation species E. globulus and E. nitens and four at-risk native eucalypts. Range-wide DNA samples of E. camaldulensis, E. cypellocarpa, E. globulus, E. nitens, E. ovata and E. viminalis, and pedigreed and putative hybrids (n = 606), were genotyped with 10 microsatellite loci. Using a two-way simulation analysis (two species in the model at a time), the accuracy of identification was 98% for first and 93% for second generation hybrids. However, the accuracy of identifying simulated backcross hybrids was lower (74%). A six-way analysis (all species in the model together) showed that as the number of species increases the accuracy of hybrid identification decreases. Despite some difficulties identifying backcrosses, the two-way Bayesian modelling approach was highly effective at identifying F1s, which, in the context of E. globulus plantations, are the primary management concern