Drivers of seedling establishment success in dryland restoration efforts.

Restoration of degraded drylands is urgently needed to mitigate climate change, reverse desertification and secure livelihoods for the two billion people who live in these areas. Bold global targets have been set for dryland restoration to restore millions of hectares of degraded land. These targets have been questioned as overly ambitious, but without a global evaluation of successes and failures it is impossible to gauge feasibility. Here we examine restoration seeding outcomes across 174 sites on six continents, encompassing 594,065 observations of 671 plant species. Our findings suggest reasons for optimism. Seeding had a positive impact on species presence: in almost a third of all treatments, 100% of species seeded were growing at first monitoring. However, dryland restoration is risky: 17% of projects failed, with no establishment of any seeded species, and consistent declines were found in seeded species as projects matured. Across projects, higher seeding rates and larger seed sizes resulted in a greater probability of recruitment, with further influences on species success including site aridity, taxonomic identity and species life form. Our findings suggest that investigations examining these predictive factors will yield more effective and informed restoration decision-making.

Eight generations of native seed cultivation reduces plant fitness relative to the wild progenitor population.

Native seed for restoration is in high demand, but widespread habitat degradation will likely prevent enough seed from being sustainably harvested from wild populations to meet this need. While propagation of native species has emerged in recent decades to address this resource gap, few studies have tested whether the processes of sampling from wild populations, followed by generations of farm cultivation, reduce plant fitness tolerance to stress over time. To test this, we grew the eighth generation of farm-propagated Clarkia pulchella Pursh (Onagraceae) alongside seeds from two of the three original wild source populations that established the native seed farm. To detect differences in stress tolerance, half of plants were subjected to a low-water treatment in the greenhouse. At the outset, farmed seeds were 4.1% heavier and had 4% greater germination compared to wild-collected seed. At maturity, farmed plants were 22% taller and had 20% larger stigmatic surfaces, even after accounting for differences in initial seed size. Importantly, the mortality of farmed plants was extremely high (75%), especially in the low-water treatment (80%). Moreover, farmed plants under the high-water treatment had 90% lower relative fitness than wild plants due to the 1.3 times greater weekly mortality and a 3-fold reduction in flowering likelihood. Together, these data suggest that bottlenecks during initial sampling and/or unconscious selection during propagation severely reduced genetic diversity and promoted inbreeding. This may undermine restoration success, especially under stressful conditions. These results indicate that more data must be collected on the effects of cultivation to determine whether it is a suitable source of restoration seed.

Managing risks related to climate variability in rangeland-based livestock production: What producer driven strategies are shared and prevalent across diverse dryland geographies?

Rangeland-based livestock production (RBLP) primarily occurs in drylands where interannual variation in rainfall directly and indirectly affects economies, plant primary productivity (forage production), and livestock reproduction and mortality. Tight ecological and economic links to climate variation constrain production in dryland systems, but producers have a breadth of strategies to reduce climate-related risks and maintain RBLP. Research on these strategies has focused on context-specific tactics linked to specific systems and/or geographies. Inspired by studies that look for broader patterns to offer frameworks for discourse and to advance collective knowledge, we review global literature to identify risk management strategies related to climate variability that are in widespread use across dryland rangeland systems and geographies. We organize strategies within three key decision areas for producers engaged in RBLP: profit and return options, land use, and herd management. Across the decision areas, four strategies emerge as playing a strong role in risk management across the globe, with refinements based on local conditions. These shared and prevalent producer driven strategies are dynamic management of forage supply (in the decision area of land use), dynamic management of animal demand (in the area of herd management), and diversification and use of social networks (both of which apply across all three decision areas). Within each of the decision areas, we found diversification reduces climate related risks but has circumstances under which it is less effective; for example, large landholders already buffered to risk via landscape diversity benefit less from livelihood diversification. In practice, implementation of the four strategies often results in livestock producers who do not maximize short-term profits but instead prioritize land resilience, large herd sizes, lifestyle goals, and longer-term economic sustainability. In this synthesis, we considered existing producer strategies for reducing risk related to climate related variability -- an intrinsic and defining characteristic of dryland rangelands -- in order to highlight valuable areas in which research can support problem solving across diverse RBLP geographies and economies, especially in a changing climate.

Decoupled recovery of ecological communities after reclamation.

Grassland restoration is largely focused on creating plant communities that match reference conditions. However, these communities reflect only a subset of the biodiversity of grassland systems. We conducted a multi-trophic study to assess ecosystem recovery following energy development for oil and gas extraction in northern US Great Plains rangelands. We compared soil factors, plant species composition and cover, and nematode trophic structuring between reclaimed oil and gas well sites ("reclaims") that comprise a chronosequence of two-33 years since reclamation and adjacent, undeveloped rangeland at distances of 50 m and 150 m from reclaim edges. Soils and plant communities in reclaims did not match those on undeveloped rangeland even after 33 years. Reclaimed soils had higher salt concentrations and pH than undeveloped soils. Reclaims had lower overall plant cover, a greater proportion of exotic and ruderal plant cover and lower native plant species richness than undeveloped rangeland. However, nematode communities appear to have recovered following reclamation. Although total and omni-carnivorous nematode abundances differed between reclaimed well sites and undeveloped rangeland, community composition and structure did not. These findings suggest that current reclamation practices recover the functional composition of nematode communities, but not soil conditions or plant communities. Our results show that plant communities have failed to recover through reclamation: high soil salinity may create a persistent impediment to native plant growth and ecosystem recovery.

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.

Oilfield Reclamation Recovers Productivity but not Composition of Arthropod Herbivores and Predators.

Arthropods are key components of grassland ecosystems. Though arthropod communities are often strongly influenced by plant communities, plants and arthropods may respond differently to disturbance. Studying plant responses alone may, therefore, not fully capture altered ecosystem dynamics; thus multi-trophic approaches are critical to fully understand ecosystem responses to disturbance. Energy development is a large-scale driver of disturbance in northern Great Plains rangelands, and recovery of arthropod communities following reclamation is not well understood. We sampled Orthoptera and spiders in western North Dakota, United States, in 2016. Samples were collected from 14 reclaimed oil well sites ('reclaims') 2-33 yr since reclamation, and native prairie at two distances (50 and 150 m) from reclaim edges. Overall Orthopteran and spider abundances on reclaims and native prairie did not differ; however, Orthopteran community composition and species abundances were distinct on reclaims versus native prairie, including increased abundances of Melanoplus femurrubrum (De Geer) (Orthoptera: Acrididae) (a noted crop pest) on reclaims. In contrast, NMS analyses revealed no differences in spider community composition between reclaims and native prairie, although abundances of one group (Salticidae) strongly decreased on reclaims. We present one of the first studies to investigate impacts of energy development and reclamation on arthropod communities. While reclamation efforts successfully recovered abundances and biomass of arthropod herbivores and predators, Orthopteran (but not spider) community composition on reclaims has not recovered to match that of intact prairie even 30 yr after reclamation. These findings suggest that energy development may have long-term or potentially irreversible impacts to rangeland arthropod communities.

Strategic plant choices can alleviate climate change impacts: A review.

Ecosystem-based adaptation (EbA) uses biodiversity and ecosystem services to reduce climate change impacts to local communities. Because plants can alleviate the abiotic and biotic stresses of climate change, purposeful plant choices could improve adaptation. However, there has been no systematic review of how plants can be applied to alleviate effects of climate change. Here we describe how plants can modify climate change effects by altering biological and physical processes. Plant effects range from increasing soil stabilization to reducing the impact of flooding and storm surges. Given the global scale of plant-related activities such as farming, landscaping, forestry, conservation, and restoration, plants can be selected strategically-i.e., planting and maintaining particular species with desired impacts-to simultaneously restore degraded ecosystems, conserve ecosystem function, and help alleviate effects of climate change. Plants are a tool for EbA that should be more broadly and strategically utilized.

Can local adaptation research in plants inform selection of native plant materials? An analysis of experimental methodologies.

Local adaptation is used as a criterion to select plant materials that will display high fitness in new environments. A large body of research has explored local adaptation in plants, however, to what extent findings can inform management decisions has not been formally evaluated. We assessed local adaptation literature for six key experimental methodologies that have the greatest effect on the application of research to selecting plant materials for natural resource management: experimental environment, response variables, maternal effects, intraspecific variation, selective agents, and spatial and temporal variability. We found that less than half of experiments used reciprocal transplants or natural field conditions, which are both informative for revegetation and restoration. Population growth rate was rarely (5%) assessed, and most studies measured only single generations (96%) and ran for less than a year. Emergence and establishment are limiting factors in successful revegetation and restoration, but the majority of studies measured later life-history stages (66%). Additionally, most studies included limited replication at the population and habitat levels and tested response to single abiotic selective factors (66%). Local adaptation research should be cautiously applied to management; future research could use alternative methodologies to allow managers to directly apply findings.

Seed production areas for the global restoration challenge.

Wild-collected seed can no longer meet global demand in restoration. Dedicated Seed Production Areas (SPA) for restoration are needed and these require application of ecological, economic, and population-genetic science. SPA design and construction must embrace the ecological sustainability principles of restoration.

Relating adaptive genetic traits to climate for Sandberg bluegrass from the intermountain western United States.

Genetic variation for potentially adaptive traits of the key restoration species Sandberg bluegrass (Poa secunda J. Presl) was assessed over the intermountain western United States in relation to source population climate. Common gardens were established at two intermountain west sites with progeny from two maternal parents from each of 130 wild populations. Data were collected over 2 years at each site on fifteen plant traits associated with production, phenology, and morphology. Analyses of variance revealed strong population differences for all plant traits (P < 0.0001), indicating genetic variation. Both the canonical correlation and linear correlation established associations between source populations and climate variability. Populations from warmer, more arid climates had generally lower dry weight, earlier phenology, and smaller, narrower leaves than those from cooler, moister climates. The first three canonical variates were regressed with climate variables resulting in significant models (P < 0.0001) used to map 12 seed zones. Of the 700 981 km(2) mapped, four seed zones represented 92% of the area in typically semi-arid and arid regions. The association of genetic variation with source climates in the intermountain west suggested climate driven natural selection and evolution. We recommend seed transfer zones and population movement guidelines to enhance adaptation and diversity for large-scale restoration projects.

Effects of Agaricus lilaceps fairy rings on soil aggregation and microbial community structure in relation to growth stimulation of western wheatgrass (Pascopyrum smithii) in Eastern Montana rangeland.

Stimulation of plant productivity caused by Agaricus fairy rings has been reported, but little is known about the effects of these fungi on soil aggregation and the microbial community structure, particularly the communities that can bind soil particles. We studied three concentric zones of Agaricus lilaceps fairy rings in Eastern Montana that stimulate western wheatgrass (Pascopyrum smithii): outside the ring (OUT), inside the ring (IN), and stimulated zone adjacent to the fungal fruiting bodies (SZ) to determine (1) soil aggregate proportion and stability, (2) the microbial community composition and the N-acetyl-ß-D-glucosaminidase activity associated with bulk soil at 0-15 cm depth, (3) the predominant culturable bacterial communities that can bind to soil adhering to wheatgrass roots, and (4) the stimulation of wheatgrass production. In bulk soil, macroaggregates (4.75-2.00 and 2.00-0.25 mm) and aggregate stability increased in SZ compared to IN and OUT. The high ratio of fungal to bacteria (fatty acid methyl ester) and N-acetyl-ß-D-glucosaminidase activity in SZ compared to IN and OUT suggest high fungal biomass. A soil sedimentation assay performed on the predominant isolates from root-adhering soil indicated more soil-binding bacteria in SZ than IN and OUT; Pseudomonas fluorescens and Stenotrophomonas maltophilia isolates predominated in SZ, whereas Bacillus spp. isolates predominated in IN and OUT. This study suggests that growth stimulation of wheatgrass in A. lilaceps fairy rings may be attributed to the activity of the fungus by enhancing soil aggregation of bulk soil at 0-15 cm depth and influencing the amount and functionality of specific predominant microbial communities in the wheatgrass root-adhering soil.

Within- and trans-generational plasticity affects the opportunity for selection in barbed goatgrass (Aegilops triuncialis).

PREMISE OF THE STUDY: Environments are composed of selective agents, and environments may also modify the efficacy of these agents. Environments affect the rate of maximum evolutionary change by influencing variation in relative fitness (i.e., the opportunity for selection, or I). Within- and transgenerational plastic environmental responses may affect I, speeding or slowing processes of local adaptation. • METHODS: We determined whether environmental factors affected the opportunity for selection (I) in Aegilops triuncialis (barbed goatgrass) by measuring I as a within- and transgenerational plastic response to two maternal glasshouse environments (serpentine/dry and loam/moist). We also determined whether this species' two most common genetic lineages (determined by DNA microsatellite length polymorphism) varied in response to glasshouse treatments. • KEY RESULTS: Opportunity for selection was less for plants grown in the dry serpentine environment than for plants grown in the moist loam environment. This response varied between genetic lineages. The east lineage exhibited a within-generation response to the dry serpentine environment. For both seed mass and average seed weight in this lineage, the opportunity for selection was lower in dry serpentine than in moist loam. The west lineage had a transgenerational response to the dry serpentine such that the opportunity for selection for seed number and seed mass was lower for plants produced by mothers grown in dry serpentine than for plants produced by mothers in moist loam. • CONCLUSIONS: Phenotypic variation in relative fitness is constrained by the dry serpentine environment, which leads to lower evolvability in this environment. Within- and transgenerational effects of the environment may slow local adaptation to serpentine soils.

Long-term population dynamics of seeded plants in invaded grasslands.

In recent decades, dozens of studies have involved attempts to introduce native and desirable nonnative plant species into grasslands dominated by invasive weeds. The newly introduced plants have proved capable of establishing, but because they are rarely monitored for more than four years, it is unknown if they have a high likelihood of persisting and suppressing invaders for the long-term. Beyond invaded grasslands, this lack of long-term monitoring is a general problem plaguing efforts to reintroduce a range of taxa into a range of ecosystems. We introduced species from seed and then periodically measured plant abundances for nine years at one site and 15 years at a second site. To our knowledge, our 15-year data are the longest to date from a seeding experiment in invaded, never-cultivated grassland. At one site, three seeded grasses maintained high densities for three or more years, but then all or nearly all individuals died. At the second site, one grass performed similarly, but two other grasses proliferated and at least one greatly suppressed the dominant invader (Centaurea maculosa). In one study, our point estimate suggests that the seeded grass Thinopyrum intermedium reduced C. maculosa biomass by 93% 15 years after seeding. In some cases, data from three and fewer years after seeding falsely suggested that seeded species were capable of persisting within the invaded grassland. In other cases, data from as late as nine years after seeding falsely suggested seeded populations would not become large enough to suppress the invader. These results show that seeded species sometimes persist and suppress invaders for long periods, but short-term data cannot predict if, when, or where this will occur. Because short-term data are not predictive of long-term seeded species performances, additional long-term data are needed to identify effective practices, traits, and species for revegetating invaded grasslands.

Native perennial grasses show evolutionary response to Bromus tectorum (cheatgrass) invasion.

Invasive species can change selective pressures on native plants by altering biotic and abiotic conditions in invaded habitats. Although invasions can lead to native species extirpation, they may also induce rapid evolutionary changes in remnant native plants. We investigated whether adult plants of five native perennial grasses exhibited trait shifts consistent with evolution in response to invasion by the introduced annual grass Bromus tectorum L. (cheatgrass), and asked how much variation there was among species and populations in the ability to grow successfully with the invader. Three hundred and twenty adult plants were collected from invaded and uninvaded communities from four locations near Reno, Nevada, USA. Each plant was divided in two and transplanted into the greenhouse. One clone was grown with B. tectorum while the other was grown alone, and we measured tolerance (ability to maintain size) and the ability to reduce size of B. tectorum for each plant. Plants from invaded populations consistently had earlier phenology than those from uninvaded populations, and in two out of four sites, invaded populations were more tolerant of B. tectorum competition than uninvaded populations. Poa secunda and one population of E. multisetus had the strongest suppressive effect on B. tectorum, and these two species were the only ones that flowered in competition with B. tectorum. Our study indicates that response to B. tectorum is a function of both location and species identity, with some, but not all, populations of native grasses showing trait shifts consistent with evolution in response to B. tectorum invasion within the Great Basin.

Patterns of introduction and adaptation during the invasion of Aegilops triuncialis (Poaceae) into Californian serpentine soils.

Multiple introductions can play a prominent role in explaining the success of biological invasions. One often cited mechanism is that multiple introductions of invasive species prevent genetic bottlenecks by parallel introductions of several distinct genotypes that, in turn, provide heritable variation necessary for local adaptation. Here, we show that the invasion of Aegilops triuncialis into California, USA, involved multiple introductions that may have facilitated invasion into serpentine habitats. Using microsatellite markers, we compared the polymorphism and genetic structure of populations of Ae. triuncialis invading serpentine soils in California to that of accessions from its native range. In a glasshouse study, we also compared phenotypic variation in phenological and fitness traits between invasive and native populations grown on loam soil and under serpentine edaphic conditions. Molecular analysis of invasive populations revealed that Californian populations cluster into three independent introductions (i.e. invasive lineages). Our glasshouse common garden experiment found that all Californian populations exhibited higher fitness under serpentine conditions. However, the three invasive lineages appear to represent independent pathways of adaptation to serpentine soil. Our results suggest that the rapid invasion of serpentine habitats in California may have been facilitated by the existence of colonizing Eurasian genotypes pre-adapted to serpentine soils.

Small variance in growth rates in annual plants has large effects on genetic drift.

PREMISE OF THE STUDY: Effective population size (N(e)) is a critical index of the evolutionary capacity of populations. Low N(e) indicates that standing genetic diversity is susceptible to loss via stochastic processes (and inbreeding) and is, therefore, unavailable for natural selection to act upon. Reported N(e) in plant populations is often quite low. What biological and ecological factors might produce such low N(e) • METHODS: We conducted a simulation model to test the effect of randomly assigned and autocorrelated growth rates of annual plants on plant-size distributions at the end of the growing season. Because plant size is directly correlated with reproductive output in annual plants, variation in plant size reflects variation in reproduction, and thus our modeled size distributions can be used to estimate N(e). • KEY RESULTS: Randomly assigned growth rates had a negligble effect on N(e)/N. Autocorrelated growth rates decreased N(e)/N as the length of the growing season increased. This was the case even when the variance in growth rates was as low as 0.1% of the mean. • CONCLUSIONS: While intrinsic plant biology can affect the degree of growth autocorrelation, ecological factors such as competition, herbivory, and abiotic stress can increase or decrease levels of growth autocorrelation. Ecological factors that increase growth autocorrelation can have significant effects on genetic drift within populations.

Coevolution between native and invasive plant competitors: implications for invasive species management.

Invasive species may establish in communities because they are better competitors than natives, but in order to remain community dominants, the competitive advantage of invasive species must be persistent. Native species that are not extirpated when highly invasive species are introduced are likely to compete with invaders. When population sizes and genetic diversity of native species are large enough, natives may be able to evolve traits that allow them to co-occur with invasive species. Native species may also evolve to become significant competitors with invasive species, and thus affect the fitness of invaders. Invasive species may respond in turn, creating either transient or continuing coevolution between competing species. In addition to demographic factors such as population size and growth rates, a number of factors including gene flow, genetic drift, the number of selection agents, encounter rates, and genetic diversity may affect the ability of native and invasive species to evolve competitive ability against one another. We discuss how these factors may differ between populations of native and invasive plants, and how this might affect their ability to respond to selection. Management actions that maintain genetic diversity in native species while reducing population sizes and genetic diversity in invasive species could promote the ability of natives to evolve improved competitive ability.

The role of adaptive trans-generational plasticity in biological invasions of plants.

High-impact biological invasions often involve establishment and spread in disturbed, high-resource patches followed by establishment and spread in biotically or abiotically stressful areas. Evolutionary change may be required for the second phase of invasion (establishment and spread in stressful areas) to occur. When species have low genetic diversity and short selection history, within-generation phenotypic plasticity is often cited as the mechanism through which spread across multiple habitat types can occur. We show that trans-generational plasticity (TGP) can result in pre-adapted progeny that exhibit traits associated with increased fitness both in high-resource patches and in stressful conditions. In the invasive sedge, Cyperus esculentus, maternal plants growing in nutrient-poor patches can place disproportional number of propagules into nutrient-rich patches. Using the invasive annual grass, Aegilops triuncialis, we show that maternal response to soil conditions can confer greater stress tolerance in seedlings in the form of greater photosynthetic efficiency. We also show TGP for a phenological shift in a low resource environment that results in greater stress tolerance in progeny. These lines of evidence suggest that the maternal environment can have profound effects on offspring success and that TGP may play a significant role in some plant invasions.