Annual-perennial lifespan variation in Chaenactis douglasii suggests a drought escape strategy in warm-arid environments.

PREMISE: Intraspecific variation in drought resistance traits, such as drought escape, appear to be frequent within wild, ruderal forb species. Understanding how these traits are arrayed across the landscape, particularly in association with climate, is critical to developing forbs for wildland restoration programs. Use of forbs is requisite for maintaining biological diversity and ecological services. METHODS: Using 6074 greenhouse-grown Chaenactis douglasii seedlings from 95 wild, seed-sourced populations across the western United States, we recorded bolting phenology and estimated genome size using flow cytometry. Mixed-effects regression models were used to assess whether climate of seed origin was predictive for bolting phenology and genome size. RESULTS: Variation in bolting, reflecting an annual vs. perennial lifespan in this species, was observed in 8.7% of the plants, with bolting plants disproportionately occurring in locations with warm, arid climates. Populations with increasing heat and aridity were positively correlated with observed bolting (r = 0.61, p < 0.0001). About one-third (22%) of the total (61%) lifespan variation was attributed to seed source climate and annual heat moisture index, a measure of aridity. Genome size had no significant effect on bolting. Projected climate modeling for mid-century (2041-2070) supports an increasing occurrence of annual lifespan. CONCLUSIONS: Our analyses support a drought escape, bet-hedging strategy in C. douglasii. Populations exposed to greater aridity exhibited a higher proportion of individuals with an annual lifespan. Drought escape leading to an annual lifespan can affect how seeds are propagated and deployed for climate-informed restoration.

Evolution of marginal populations of an invasive vine increases the likelihood of future spread.

The prediction of invasion patterns may require an understanding of intraspecific differentiation in invasive species and its interaction with climate change. We compare Japanese honeysuckle (Lonicera japonica) plants from the core (100-150 yr old) and northern margin (< 65 yr old) of their North American invaded range to determine whether evolution during invasion increases the probability of future expansion. Plants from populations in the core and margin were compared in two sites beyond the northern range edge to assess their potential to invade novel areas. Data were compared with previous work to assess the effect of latitudinal climate on L. japonica spread. Winter survival in current climates was modeled and projected for future climates to predict future spread. Margin plants were larger and had 60% greater survival than core plants at sites beyond the northern range edge. Overall, winter survival decreased with increasing latitude and decreasing temperature, and was greater in margin plants than core plants. Models suggested that greater winter tolerance in margin populations has increased L. japonica's northward spread by 76 km, and that this survival advantage will persist under future climates. These results demonstrate that evolution during invasion may increase spread beyond predictions using increasing global temperatures alone.

Strong patterns of intraspecific variation and local adaptation in Great Basin plants revealed through a review of 75 years of experiments.

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.

Population history provides foundational knowledge for utilizing and developing native plant restoration materials.

A species' population structure and history are critical pieces of information that can help guide the use of available native plant materials in restoration treatments and decide what new native plant materials should be developed to meet future restoration needs. In the western United States, Pseudoroegneria spicata (bluebunch wheatgrass; Poaceae) is an important component of grassland and shrubland plant communities and commonly used for restoration due to its drought resistance and competitiveness with exotic weeds. We used next-generation sequencing data to investigate the processes that shaped P. spicata's geographic pattern of genetic variation across the Intermountain West. Pseudoroegneria spicata's genetic diversity is partitioned into populations that likely differentiated since the Last Glacial Maximum. Adjacent populations display varying magnitudes of historical gene flow, with migration rates ranging from multiple migrants per generation to multiple generations per migrant. When considering the commercial germplasm sources available for restoration, genetic identities remain representative of the wildland localities from which germplasm sources were originally developed, and they maintain high levels of heterozygosity and nucleotide diversity. However, the commercial germplasm sources represent a small fraction of the overall genetic diversity of P. spicata in the Intermountain West. Given the low migration rates and long divergence times between some pairs of P. spicata populations, using commercial germplasm sources could facilitate undesirable restoration outcomes when used in certain geographic areas, even if the environment in which the commercial materials thrive is similar to that of the restoration site. As such, population structure and history can be used to provide guidance on what geographic areas may need additional native plant materials so that restoration efforts support species and community resilience and improve outcomes.

Adaptive divergence at the margin of an invaded range.

Invasive plant species threaten biological communities globally. However, relatively little is known about how evolutionary processes vary over the course of an invasion. To evaluate the importance of historical and adaptive drivers of range expansion, we compare the performance of North American populations of invasive Lonicera japonica from areas established 100-150 years ago, now the southern core of the range, to populations from the northern range margin, established within the last 65 years. Growth and survival of individuals from 17 core and 14 margin populations were compared in common gardens at both regions. After three years, margin plants were larger than core plants regardless of planting region, with 34% more branches and 36% greater biomass. Growth rate was directly related to survival, and margin plants also had 30% greater survival than core plants across both regions. Larger size of individuals from margin populations suggests either that the shorter growing period at the northern margin has selected for more rapid growth or that range expansion has selected for plants with a greater colonizing ability, including rapid establishment and growth. Because this evolution has resulted in enhanced survival and increased growth rate it may drive spread, increasing the likelihood of further invasion.

Genetic variation in adaptive traits and seed transfer zones for Pseudoroegneria spicata (bluebunch wheatgrass) in the northwestern United States.

A genecological approach was used to explore genetic variation in adaptive traits in Pseudoroegneria spicata, a key restoration grass, in the intermountain western United States. Common garden experiments were established at three contrasting sites with seedlings from two maternal parents from each of 114 populations along with five commercial releases commonly used in restoration. Traits associated with size, flowering phenology, and leaf width varied considerably among populations and were moderately correlated with the climates of the seed sources. Pseudoroegneria spicata populations from warm, arid source environments were smaller with earlier phenology and had relatively narrow leaves than those from mild climates with cool summers, warm winters, low seasonal temperature differentials, high precipitation, and low aridity. Later phenology was generally associated with populations from colder climates. Releases were larger and more fecund than most of the native ecotypes, but were similar to native populations near their source of origin. Differences among native populations associated with source climates that are logical for survival, growth, and reproduction indicate that genetic variation across the landscape is adaptive and should be considered during restoration. Results were used to delineate seed transfer zones and population movement guidelines to ensure adapted plant materials for restoration activities.

Reproductive success in varying light environments: direct and indirect effects of light on plants and pollinators.

Plant populations often exist in spatially heterogeneous environments. Light level can directly affect plant reproductive success through resource availability or by altering pollinator behavior. It can also indirectly influence reproductive success by determining floral display size which may in turn influence pollinator attraction. We evaluated direct and indirect effects of light availability and measured phenotypic selection on phenological traits that may enhance pollen receipt in the insect-pollinated herb Campanulastrum americanum. In a natural population, plants in the sun had larger displays and received 7 times more visits than plants in the shade. Using experimental arrays to separate the direct effects of irradiance on insects from their response to display size, we found more visits to plants in the sun than in the shade, but no association between number of visits each flower received and display size. Plants in the sun were not pollen limited but pollen-augmented shade flowers produced 50% more seeds than open-pollinated flowers. Phenological traits, which may influence pollen receipt, were not under direct selection in the sun. However, earlier initiation and a longer duration of flowering were favored in the shade, which may enhance visitation in this pollen-limited habitat.