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            Zhou, Jun (Ed.)IntroductionClimate 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. MethodsWe 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. ResultsFour 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. DiscussionResults 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.more » « less
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            Abstract Understanding how vegetation responds to drought is fundamental for understanding the broader implications of climate change on foundation tree species that support high biodiversity. Leveraging remote sensing technology provides a unique vantage point to explore these responses across and within species.We investigated interspecific drought responses of twoPopulusspecies (P.fremontii,P.angustifolia) and their naturally occurring hybrids using leaf‐level visible through shortwave infrared (VSWIR; 400–2500 nm) reflectance. AsF1hybrids backcross with either species, resulting in a range of backcross genotypes, we heretofore refer to the two species and their hybrids collectively as ‘cross types’. We additionally explored intraspecific variation inP. fremontiidrought response at the leaf and canopy levels using reflectance data and thermal unmanned aerial vehicle (UAV) imagery. We employed several analyses to assess genotype‐by‐environment (G × E) interactions concerning drought, including principal component analysis, support vector machine and spectral similarity index.Five key findings emerged: (1) Spectra of all three cross types shifted significantly in response to drought. The magnitude of these reaction norms can be ranked from hybrids>P. fremontii>P. angustifolia, suggesting differential variation in response to drought; (2) Spectral space among cross types constricted under drought, indicating spectral—and phenotypic—convergence; (3) Experimentally, populations ofP. fremontiifrom cool regions had different responses to drought than populations from warm regions, with source population mean annual temperature driving the magnitude and direction of change in VSWIR reflectance. (4) UAV thermal imagery revealed that watered, warm‐adapted populations maintained lower leaf temperatures and retained more leaves than cool‐adapted populations, but differences in leaf retention decreased when droughted. (5) These findings are consistent with patterns of local adaptation to drought and temperature stress, demonstrating the ability of leaf spectra to detect ecological and evolutionary responses to drought as a function of adaptation to different environments.Synthesis.Leaf‐level spectroscopy and canopy‐level UAV thermal data captured inter‐ and intraspecific responses to water stress in cottonwoods, which are widely distributed in arid environments. This study demonstrates the potential of remote sensing to monitor and predict the impacts of drought on scales varying from leaves to landscapes.more » « less
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            ABSTRACT 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.more » « less
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            Summary Populus fremontiiis among the most dominant, and ecologically important riparian tree species in the western United States and can thrive in hyper‐arid riparian corridors. Yet,P. fremontiiforests have rapidly declined over the last decade, particularly in places where temperatures sometimes exceed 50°C.We evaluated high temperature tolerance of leaf metabolism, leaf thermoregulation, and leaf hydraulic function in eightP. fremontiipopulations spanning a 5.3°C mean annual temperature gradient in a well‐watered common garden, and at source locations throughout the lower Colorado River Basin.Two major results emerged. First, despite having an exceptionally highTcrit(the temperature at which Photosystem II is disrupted) relative to other tree taxa, recent heat waves exceededTcrit, requiring evaporative leaf cooling to maintain leaf‐to‐air thermal safety margins. Second, in midsummer, genotypes from the warmest locations maintained lower midday leaf temperatures, a higher midday stomatal conductance, and maintained turgor pressure at lower water potentials than genotypes from more temperate locations.Taken together, results suggest that under well‐watered conditions,P. fremontiican regulate leaf temperature belowTcritalong the warm edge of its distribution. Nevertheless, reduced Colorado River flows threaten to lower water tables below levels needed for evaporative cooling during episodic heat waves.more » « less
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            Abstract 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 asPopulus, 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 leaf nitrogen. (2) For both defense and structural compounds, plastic responses to different garden climates depended on the climate in which a population or genotype originated. Specifically, trees originating from cool provenances showed higher defense plasticity in response to climate changes than trees from warmer provenances. (3) Trees from cool provenances growing in cool garden conditions expressed the lowest constitutive defense levels but the strongest induced (plastic) defenses in response to damage. (4) The combination of hot garden conditions and simulated herbivory switched the strategy used by these genotypes, increasing constitutive defenses but erasing the capacity for induction after damage. Because Fremont cottonwood chemistry plays a major role in shaping riparian communities and ecosystems, 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.more » « less
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            Increasing heatwaves are threatening forest ecosystems globally. Leaf thermal regulation and tolerance are important for plant survival during heatwaves, though the interaction between these processes and water availability is unclear. Genotypes of the widely distributed foundation tree speciesPopulus fremontiiwere studied in a controlled common garden during a record summer heatwave—where air temperature exceeded 48 °C. When water was not limiting, all genotypes cooled leaves 2 to 5 °C below air temperatures. Homeothermic cooling was disrupted for weeks following a 72-h reduction in soil water, resulting in leaf temperatures rising 3 °C above air temperature and 1.3 °C above leaf thresholds for physiological damage, despite the water stress having little effect on leaf water potentials. Tradeoffs between leaf thermal safety and hydraulic safety emerged but, regardless of water use strategy, all genotypes experienced significant leaf mortality following water stress. Genotypes from warmer climates showed greater leaf cooling and less leaf mortality after water stress in comparison with genotypes from cooler climates. These results illustrate how brief soil water limitation disrupts leaf thermal regulation and potentially compromises plant survival during extreme heatwaves, thus providing insight into future scenarios in which ecosystems will be challenged with extreme heat and unreliable soil water access.more » « less
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            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 experi- mental 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 adjust- ments. 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.more » « less
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            Mycorrhizal restoration benefits are widely acknowledged, yet factors underpinning this success remain unclear. To illuminate when natural regeneration might be sufficient, we investigated the degree mycorrhizal fungi would colonizePopulus fremontii(Fremont cottonwood) 2 years after the restoration of a riparian corridor, in the presence of an adjacent source. We compared colonization levels across plant populations and ecotypes, and from trees in the planted area to those in natural source populations. Four findings contribute to the theory and application of host–symbiont interactions. (1) Median ectomycorrhizal colonization of trees in the planted area was less than one‐tenth of that within natural source populations (p < 0.05), suggesting that even with adjacent intact habitat, sluggish regeneration would make proactive mycorrhizal restoration beneficial. (2) Within the planted area, median ectomycorrhizal and arbuscule colonization of trees sourced from greater distances were less than one‐third of that for trees sourced locally (p < 0.05), suggesting translocation poses barriers to symbioses. (3) Changes in colonization did not align with plant ecotypes, suggesting that geographic scales of selection for plants and fungi differ. (4) Slight increases in median mycorrhizal colonization (from 0% to 5%) were strongly correlated with increased survival for the plant provenance with lowest survival (r2 = 46% andrs = 48%,p < 0.05), suggesting mycorrhizae are particularly beneficial when plants are under stress (including translocation‐induced stress). This study is novel in demonstrating that mycorrhizal regeneration is slow even in the presence of adjacent intact habitat, and that when colonization could seem negligible, it may still have biological significance.more » « less
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            Ahmed, Ferdous (Ed.)We addressed the hypothesis that intraspecific genetic variation in plant traits from different sites along a distance/elevation gradient would influence the communities they support when grown at a new site. Answers to this hypothesis are important when considering the community consequences of assisted migration under climate change; i.e., if you build it will they come?. We surveyed arthropod communities occurring on the foundation riparian tree species Populus angustifolia along a distance/elevation gradient and in a common garden where trees from along the gradient were planted 20–22 years earlier. Three major patterns were found: 1) In the wild, arthropod community composition changed significantly. Trees at the lower elevation site supported up to 58% greater arthropod abundance and 26% greater species richness than more distant, high elevation trees. 2) Trees grown in a common garden sourced from the same locations along the gradient, supported arthropod communities more similar to their corresponding wild trees, but the similarity declined with transfer distance and elevation. 3) Of five functional traits examined, leaf area, a trait under genetic control that decreases at higher elevations, is correlated with differences in arthropod species richness and abundance. Our results suggest that genetic differences in functional traits are stronger drivers of arthropod community composition than phenotypic plasticity of plant traits due to environmental factors. We also show that variation in leaf area is maintained and has similar effects at the community level while controlling for environment. These results demonstrate how genetically based traits vary across natural gradients and have community-level effects that are maintained, in part, when they are used in assisted migration. Furthermore, optimal transfer distances for plants suffering from climate change may not be the same as optimal transfer distances for their dependent communities.more » « less
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