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Leaf Economic and Hydraulic Traits Signal Disparate Climate Adaptation Patterns in Two Co-Occurring Woodland Eucalypts
Publication . Prober, Suzanne M.; Potts, Brad M.; Harrison, Peter A.; Wiehl, Georg; Bailey, Tanya G.; Costa e Silva, João; Price, Meridy R.; Speijers, Jane; Steane, Dorothy A.; Vaillancourt, René E.
With climate change impacting trees worldwide, enhancing adaptation capacity has become
an important goal of provenance translocation strategies for forestry, ecological renovation, and
biodiversity conservation. Given that not every species can be studied in detail, it is important
to understand the extent to which climate adaptation patterns can be generalised across species,
in terms of the selective agents and traits involved. We here compare patterns of genetic-based
population (co)variation in leaf economic and hydraulic traits, climate–trait associations, and genomic
differentiation of two widespread tree species (Eucalyptus pauciflora and E. ovata). We studied 2-yearold
trees growing in a common-garden trial established with progeny from populations of both
species, pair-sampled from 22 localities across their overlapping native distribution in Tasmania,
Australia. Despite originating from the same climatic gradients, the species differed in their levels of
population variance and trait covariance, patterns of population variation within each species were
uncorrelated, and the species had different climate–trait associations. Further, the pattern of genomic
differentiation among populations was uncorrelated between species, and population differentiation
in leaf traits was mostly uncorrelated with genomic differentiation. We discuss hypotheses to
explain this decoupling of patterns and propose that the choice of seed provenances for climatebased
plantings needs to account for multiple dimensions of climate change unless species-specific
information is available
Linking leaf economic and hydraulic traits with early-age growth performance and survival of Eucalyptus pauciflora
Publication . Costa e Silva, João; Potts, Brad M.; Wiehl, Georg; Prober, Suzanne M.
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 environment
Directional Selection on Tree Seedling Traits Driven by Experimental Drought Differs Between Mesic and Dry Populations
Publication . Costa e Silva, João; Jordan, Rebecca; Potts, Brad M.; Pinkard, Elisabeth; Prober, Suzanne M.
We 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 traits
Performance-based inference of selection on stomatal length and specific leaf area varies with climate-of-origin of the forest tree, Eucalyptus ovata
Publication . Costa e Silva, João; Potts, Brad M.; Prober, Suzanne M.
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.
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Funding agency
Fundação para a Ciência e a Tecnologia
Funding programme
DL 57/2016
Funding Award Number
DL 57/2016/CP1382/CT0008