Variation in natural selection across heterogeneous landscapes often produces (a) among‐population differences in phenotypic traits, (b) trait‐by‐environment associations, and (c) higher fitness of local populations. Using a broad literature review of common garden studies published between 1941 and 2017, we documented the commonness of these three signatures in plants native to North America's Great Basin, an area of extensive restoration and revegetation efforts, and asked which traits and environmental variables were involved. We also asked, independent of geographic distance, whether populations from more similar environments had more similar traits. From 327 experiments testing 121 taxa in 170 studies, we found 95.1% of 305 experiments reported among‐population differences, and 81.4% of 161 experiments reported trait‐by‐environment associations. Locals showed greater survival in 67% of 24 reciprocal experiments that reported survival, and higher fitness in 90% of 10 reciprocal experiments that reported reproductive output. A meta‐analysis on a subset of studies found that variation in eight commonly measured traits was associated with mean annual precipitation and mean annual temperature at the source location, with notably strong relationships for flowering phenology, leaf size, and survival, among others. Although the Great Basin is sometimes perceived as a region of homogeneous ecosystems, our results demonstrate widespread habitat‐related population differentiation and local adaptation. Locally sourced plants likely harbor adaptations at rates and magnitudes that are immediately relevant to restoration success, and our results suggest that certain key traits and environmental variables should be prioritized in future assessments of plants in this region.
Partnerships between researchers and restoration practitioners can improve restoration outcomes, which is especially important for restoration in challenging settings. Here, we describe one such partnership in the Great Basin, United States, which used trait-based methods and practitioner knowledge to identify the most promising seed sources for restoration. Managers in this region can either use widely available commercial seeds, which are often sourced from far outside the seeding area, or take extra steps to use locally collected seeds. We asked whether local, wild-collected seeds of two native plants, Elymus elymoides and Poa secunda, had traits more conducive to seedling establishment in degraded sites, relative to commercial sources. Seeds were collected from four remnant native populations within lands managed by the Winnemucca Bureau of Land Management. Collections were screened for seed and seedling characteristics previously identified as associated with increased seedling performance in degraded Great Basin systems, and we provide a detailed methodology for these measurements. Relative to commercial seeds, wild-collected seeds had more characteristics identified as beneficial for seedling establishment, including phenology, root allocation, root form, and overall size characteristics that suggest locally sourced populations would be likely to establish better than commercial sources. Using phenotypic traits as criteria, the most promising wild populations were selected for agronomic production to increase the quantities of seeds available for restoration, and field trials are ongoing using these field-increased seeds. These results provide support for collaborative efforts to identify, collect, screen, and increase the availability of local seed sources to improve restoration success.
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The spatial structure of genomic and phenotypic variation across populations reflects historical and demographic processes as well as evolution via natural selection. Characterizing such variation can provide an important perspective for understanding the evolutionary consequences of changing climate and for guiding ecological restoration. While evidence for local adaptation has been traditionally evaluated using phenotypic data, modern methods for generating and analyzing landscape genomic data can directly quantify local adaptation by associating allelic variation with environmental variation. Here, we analyze both genomic and phenotypic variation of rubber rabbitbrush (Ericameria nauseosa), a foundational shrub species of western North America. To quantify landscape genomic structure and provide perspective on patterns of local adaptation, we generated reduced representation sequencing data for 17 wild populations (222 individuals; 38,615 loci) spanning a range of environmental conditions. Population genetic analyses illustrated pronounced landscape genomic structure jointly shaped by geography and environment. Genetic-environment association (GEA) analyses using both redundancy analysis (RDA) and a machine-learning approach (Gradient Forest) indicated environmental variables (precipitation seasonality, slope, aspect, elevation, and annual precipitation) influenced spatial genomic structure, and were correlated with allele frequency shifts indicative of local adaptation at a consistent set of genomic regions. We compared our GEA based inference of local adaptation with phenotypic data collected by growing seeds from each population in a greenhouse common garden. Population differentiation in seed weight, emergence, and seedling traits was associated with environmental variables (e.g., precipitation seasonality) that were also implicated in GEA analyses, suggesting complementary conclusions about the drivers of local adaptation across different methods and data sources. Our results provide a baseline understanding of spatial genomic structure for E. nauseosa across the western Great Basin and illustrate the utility of GEA analyses for detecting the environmental causes and genetic signatures of local adaptation in a widely distributed plant species of restoration significance.
The UN Decade on Ecosystem Restoration will result in an unprecedented need for seeds. Agricultural production, or the growing of plants under controlled conditions to produce desired resources, can be a helpful tool for providing the quantities of seeds needed for large‐scale restoration. In some ecosystems, agricultural production of native plant seeds is part of the restoration process. This is particularly true for native plants that are similar to major food crops, such as native grasses. However, conventional crops have contrasting characteristics (e.g. fast growing, high yield for a focused use) from those desired in restored vegetation (plants capable of surviving and providing ecosystem services in variable environments). This contrast leads to significant, often opposing differences in desirable characteristics in crop versus natural systems. Examples include rapid germination in domesticated crops versus seed dormancy in wild plants, uniform germination, growth and flowering versus bet‐hedging strategies, or high seed retention versus seed dispersal. While it may be useful to maximize productivity of native plants in agricultural environments with similar trait modifications, for restoration, the demands of the natural environment should be primary, and agricultural production should aim to maximize the diversity, function, and evolutionary potential of native species. Any perceived benefits that come from the production of large quantities of inexpensive seeds that cannot meet the needs of wild communities are short‐sighted. We suggest research directions that will be needed to meld agricultural production needs with those of restoration practitioners, as a challenge for those invested in the procurement and use of wild seeds.
Trait–environment correlations can arise from local adaptation and can identify genetically and environmentally appropriate seeds for restoration projects. However, anthropogenic changes can disrupt the relationships between traits and fitness. Finding the best seed sources for restoration may rely on describing plant traits adaptive in disturbed and invaded environments, recognizing that while traits may differ among species and functional groups, there may be similarities in the strategies that increase seedling establishment. Focusing on three grass genera, two shrub species, and two forb genera, we collected seeds of all taxa from 16 common sites in the sagebrush steppe of the western United States. We measured seed and seedling characteristics, including seed size, emergence timing, and root and shoot traits, and compiled a suite of environmental variables for each collection site. We described trait–environment associations and asked how traits or environment of origin were associated with seedling survival in invaded gardens. Sampling seven taxa from the same sites allowed us to ask how trait–environment–performance associations differ among taxa and whether natural selection favors similar traits across multiple taxa and functional groups. All taxa showed trait–environment associations consistent with local adaptation, and both environment of origin and phenotypes predicted survival in competitive restoration settings, with some commonalities among taxa. Notably, rapid emergence and larger seeds increased survival for multiple taxa. Environmental factors at collection sites, including lower slopes (especially for grasses), greater mean annual temperatures (especially for shrubs and forbs), and greater precipitation seasonality were frequently associated with increased survival. We noted one collection site with high seedling survival across all seven taxa, suggesting that conditions within some sites may result in selection for traits that increase establishment for multiple species. Thus, choosing native plant sources with the most adaptive traits, along with matching climates, will likely improve the restoration of invaded communities.
Trait-environment correlations can arise from local adaptation and can identify genetically and environmentally appropriate seeds for restoration projects. However, anthropogenic changes can disrupt the relationships between traits and fitness. Finding the best seed sources for restoration may rely on describing plant traits adaptive in disturbed and invaded environments, recognizing that while traits may differ among species and functional groups, there may be similarities in the strategies that increase seedling establishment. Focusing on three grass genera, two shrub species, and two forb genera, we collected seeds of all taxa from 16 common sites in the sagebrush steppe of the western United States. We measured seed and seedling characteristics, including seed size, emergence timing, and root and shoot traits, and compiled a suite of environmental variables for each collection site. We described trait-environment associations and asked how traits or environment of origin were associated with seedling survival in invaded gardens. Sampling seven taxa from the same sites allowed us to ask how trait-environment-performance associations differ among taxa and whether natural selection favors similar traits across multiple taxa and functional groups. All taxa showed trait-environment associations consistent with local adaptation, and both environment of origin and phenotypic traits predicted survival in competitive restoration settings, with some commonalities among taxa. Notably, rapid emergence and larger seeds increased survival for multiple taxa. Environmental factors at collection sites, including lower slopes (especially for grasses), greater mean annual temperatures (especially for shrubs and forbs), and greater precipitation seasonality were frequently associated with increased survival. We noted one collection site with high seedling survival across all seven taxa, suggesting that conditions within some sites may result in selection for traits that increase establishment for multiple species. Thus, choosing native plant sources with the most adaptive traits, along with matching climates, will likely improve the restoration of invaded communities.
Establishing plants from seed is often a limitation to restoration success in semi-arid systems.For restoration purposes, managers can either use widely-available commercial seeds, which are often sourced from far outside the seeding area, or take extra steps to use locally collected seeds.If local seeds have traits more conducive to seedling establishment in degraded sites, they could increase restoration success. Here, we asked whether wild-collected seeds of two native perennial grasses, Elymus elymoides and Poa secunda, had more favorable characteristics than commercial sources. Seeds were collected from four populations within the Winnemucca District of the Nevada Bureau of Land Management, which manages lands within the Great Basin, US.Collections were screened for seed and seedling characteristics associated with increased plant performance in invaded Great Basin systems, and we provide a detailed methodology for these measurements. Relative to commercial seeds, wild-collected seeds had more characteristics identified as beneficial for seedling establishment including earlier emergence, higher specific root length, more root tips, and smaller overall size (E. elymoides), and earlier emergence, longer roots, higher root mass ratio, and more root tips (P. secunda). Commercial sources had significantly larger seeds than wild populations, a trait that had mixed effects on performance in previous research, and one that may change as wild collections are increased in agronomic conditions. These results suggest that locally-sourced populations are more likely to perform well in invaded areas, providing support for efforts to collect, screen, and increase local sources of seeds to improve restoration success. Author contributions:EL, SB, OB, RB, and MW conceived and designed the research; SB and AA performed the experiments; EL analyzed the data; all authors contributed to writing and editing the manuscript. Running head Seedling traits for restoration Implications for Practice• Collecting and increasing local seeds requires more time and effort than purchasing commercially-available seeds, but if these sources have a greater chance at surviving in restoration sites, this effort is warranted • In our study, remnant local populations possessed more potentially adaptive traits than commercially-available alternatives, indicating they may be superior sources for the restoration of disturbed sites in their local regions • Screening multiple seed sources for potentially adaptive seed and seedling traits can be a relatively quick and effective way to select the most promising seeds for increase
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