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Climate means and variability are shifting rapidly, leading to mismatches between climate and locally adapted plant traits. Phenotypic plasticity, the ability of a plant to respond to environmental conditions within a lifetime, may provide a buffer for plants to persist under increasing temperature and water stress. We used two reciprocal common gardens across a steep temperature gradient to investigate plasticity in six populations of Fremont cottonwood, an important foundation tree species in arid riparian ecosystems. We investigated two components of leaf hydraulic architecture: Leaf venation and stomatal morphology, both of which regulate leaf water potential and photosynthesis. These traits will likely affect plant performance under climate stressors, but it is unclear whether they are controlled by genetic or environmental factors and whether they respond to the environment in parallel or independent directions. We found that: (1) Populations had divergent responses to a hotter growing environment, increasing or decreasing vein density. (2) Populations showed surprisingly independent responses of venation vs. stomatal traits. (3) As a result of these different responses, plasticity in hydraulic architecture traits was not predictable from historic climate conditions at population source locations and often varied substantially within populations. (4) Hydraulic architecture was clearly linked to growth, with higher vein and stomatal density predicting greater tree growth in the hottest growing environment. However, higher plasticity in these traits did not increase average growth across multiple environments. Thus,P. fremontiipopulations and genotypes vary in their capacity to adjust their leaf hydraulic architecture and support growth in contrasting environments, but this plasticity is not clearly predictable or beneficial. Identifying genotypes suitable for future conditions will depend on the relative importance of multiple traits and on both evolutionary and ecological responses to changing temperature and water availability.

 

Plants respond to rapid environmental change in ways that depend on both their genetic identity and their phenotypic plasticity, impacting their survival as well as associated ecosystems. However, genetic and environmental effects on phenotype are difficult to quantify across large spatial scales and through time. Leaf hyperspectral reflectance offers a potentially robust approach to map these effects from local to landscape levels. Using a handheld field spectrometer, we analyzed leaf‐level hyperspectral reflectance of the foundation tree species Populus fremontii in wild populations and in three 6‐year‐old experimental common gardens spanning a steep climatic gradient. First, we show that genetic variation among populations and among clonal genotypes is detectable with leaf spectra, using both multivariate and univariate approaches. Spectra predicted population identity with 100% accuracy among trees in the wild, 87%–98% accuracy within a common garden, and 86% accuracy across different environments. Multiple spectral indices of plant health had significant heritability, with genotype accounting for 10%–23% of spectral variation within populations and 14%–48% of the variation across all populations. Second, we found gene by environment interactions leading to population‐specific shifts in the spectral phenotype across common garden environments. Spectral indices indicate that genetically divergent populations made unique adjustments to their chlorophyll and water content in response to the same environmental stresses, so that detecting genetic identity is critical to predicting tree response to change. Third, spectral indicators of greenness and photosynthetic efficiency decreased when populations were transferred to growing environments with higher mean annual maximum temperatures relative to home conditions. This result suggests altered physiological strategies further from the conditions to which plants are locally adapted. Transfers to cooler environments had fewer negative effects, demonstrating that plant spectra show directionality in plant performance adjustments. Thus, leaf reflectance data can detect both local adaptation and plastic shifts in plant physiology, informing strategic restoration and conservation decisions by enabling high resolution tracking of genetic and phenotypic changes in response to climate change. 

Climate change has increased the need for forest restoration, but low planting success and limited availability of planting materials hamper these efforts. Invasive plants and their soil legacies can further reduce restoration success. Thus, strategies that optimize restoration are crucial. Assisted migration and inoculation with native microbial symbiont communities have great potential to increase restoration success. However, assisted migrants can still show reduced survival compared to local provenances depending on transfer distance. Inoculation with mycorrhizal fungi, effective if well-matched to plants and site conditions, can have neutral to negative results with poor pairings. Few studies have examined the interaction between these two strategies in realistic field environments where native plants experience the combined effects of soil legacies left by invasive plants and the drought conditions that result from a warming, drying climate.

We planted two ecotypes (local climate and warmer climate) ofPopulus fremontii(Fremont cottonwoods), in soils with and without legacies of invasion byTamarixspp. (tamarisk), and with and without addition of native mycorrhizal fungi and other soil biota from the warmer climate.

Four main results emerged. 1) First year survival in soil legacies left behind after tamarisk invasion and removal was less than one tenth of survival in soil without a tamarisk legacy. 2) Actively restoring soil communities after tamarisk removal tripled first year cottonwood survival for both ecotypes, but only improved survival of the warmer, assisted migrant ecotype trees in year two. 3) Actively restoring soil communities in areas without a tamarisk history reduced first year survival for both ecotypes, but improved survival of the warmer, assisted migrant ecotype trees in year two. 4) By the second year, inoculated assisted migrants survived at five times the rate of inoculated trees from the local ecotype.

Results emphasize the detrimental effects of soil legacies left after tamarisk invasion and removal, the efficacy of assisted migration and restoring soil communities alongside plants, and the need to thoughtfully optimize pairings between plants, fungi, and site conditions.

 

Leaf-level reflectance data from multiple genotypes nested within multiple populations each grown in 2-3 common gardens. Both population provenances and common garden locations span the climate gradient of Arizona, USA. 

Despite an increased focus on multiscale relationships and interdisciplinary integration, few macroecological studies consider the contribution of genetic-based processes to landscape-scale patterns. We test the hypothesis that tree genetics, climate, and geography jointly drive continental-scale patterns of community structure, using genome-wide SNP data from a broadly distributed foundation tree species (Populus fremontii S. Watson) and two dependent communities (leaf-modifying arthropods and fungal endophytes) spanning southwestern North America. Four key findings emerged: (1) Tree genetic structure was a significant predictor for both communities; however, the strength of influence was both scale- and community-dependent. (2) Tree genetics was the primary driver for endophytes, explaining 17% of variation in continental-scale community structure, whereas (3) climate was the strongest predictor of arthropod structure (24%). (4) Power to detect tree genotype—community phenotype associations changed with scale of genetic organization, increasing from individuals to populations to ecotypes, emphasizing the need to consider nonstationarity (i.e., changes in the effects of factors on ecological processes across scales) when inferring macrosystem properties. Our findings highlight the role of foundation tree species as drivers of macroscale community structure and provide macrosystems ecology with a theoretical framework for linking fine- and intermediate-scale genetic processes to landscape-scale patterns. Management of the genetic diversity harbored within foundation species is a critical consideration for conserving and sustaining regional biodiversity. 

Selection on quantitative traits by heterogeneous climatic conditions can lead to substantial trait variation across a species range. In the context of rapidly changing environments, however, it is equally important to understand selection on trait plasticity. To evaluate the role of selection in driving divergences in traits and their associated plasticities within a widespread species, we compared molecular and quantitative trait variation in Populus fremontii (Fremont cottonwood), a foundation riparian distributed throughout Arizona. Using SNP data and genotypes from 16 populations reciprocally planted in three common gardens, we first performed QST-FST analyses to detect selection on traits and trait plasticity. We then explored the environmental drivers of selection using trait-climate and plasticity-climate regressions. Three major findings emerged: 1) There was significant genetic variation in traits expressed in each of the common gardens and in the phenotypic plasticity of traits across gardens, both of which were heritable. 2) Based on QST-FST comparisons, there was evidence of selection in all traits measured; however, this result varied from no effect in one garden to highly significant in another, indicating that detection of past selection is environmentally dependent. We also found strong evidence of divergent selection on plasticity across environments for two traits. 3) Traits and/or their plasticity were often correlated with population source climate (R2 up to 0.77 and 0.66, respectively). These results suggest that steep climate gradients across the Southwest have played a major role in shaping the evolution of divergent phenotypic responses in populations and genotypes now experiencing climate change. 

Climate change is threatening the persistence of many tree species via independent and interactive effects on abiotic and biotic conditions. In addition, changes in temperature, precipitation, and insect attacks can alter the traits of these trees, disrupting communities and ecosystems. For foundation species such as Populus, phytochemical traits are key mechanisms linking trees with their environment and are likely jointly determined by interactive effects of genetic divergence and variable environments throughout their geographic range. Using reciprocal Fremont cottonwood (Populus fremontii) common gardens along a steep climatic gradient, we explored how environment (garden climate and simulated herbivore damage) and genetics (tree provenance and genotype) affect both foliar chemical traits and the plasticity of these traits. We found that: 1) Constitutive and plastic chemical responses to changes in garden climate and damage varied among defense compounds, structural compounds and nitrogen. 2) For both defense and structural compounds, plastic responses to garden climate depended on the climate in which a population or genotype evolved. Specifically, trees originating from cool provenances showed higher defense plasticity in response to climate changes than trees from hotter provenances. 3) Trees from cool provenances growing in cool conditions expressed the lowest constitutive defense levels but the strongest induced (plastic) defenses. 4) The combination of hot growing conditions and simulated herbivory switched the strategy used by these genotypes, increasing constitutive defenses but erasing the capacity for induction. Because Fremont cottonwood chemistry plays a major role in shaping riparian communities and ecosystems in the southwestern US, the effects of changes in phytochemical traits can be wide-reaching. As the southwestern US is confronted with warming temperatures and insect outbreaks, these results improve our capacity to predict ecosystem consequences of climate change and inform selection of tree genotypes for conservation and restoration purposes.