Elizabeth A. Leger

THE ADAPTIVE VALUE OF REMNANT NATIVE PLANTS IN INVADED COMMUNITIES: AN EXAMPLE FROM THE GREAT BASIN

Changes in the species composition of biotic communities may alter patterns of natural selection occurring within them. Native perennial grass species in the Intermountain West are experiencing a shift in the composition of interspecific competitors from primarily perennial species to an exotic, annual grass. Thus traits that confer an advantage to perennial grasses in the presence of novel annual competitors may evolve in invaded communities. Here I show that such traits are apparent in populations of a native perennial grass, big squirreltail (Elymus multisetus M.E. Jones), exposed to cheatgrass (Bromus tectorum L.) competitors. Dormant big squirreltail plants were collected from cheatgrass-invaded and uninvaded sites near Bordertown, California, USA, a mid-elevation (1600 m) sagebrush community, and transplanted into pots in a greenhouse. Individual plants were split into equal halves. One half was grown with competition from cheatgrass, and the other half was grown without competition. Plants collected from invaded sites responded more quickly to watering, growing more leaves in the first 10 days after transplanting. In addition, big squirreltail plants collected from invaded areas experienced a smaller decrease in plant size when grown with competition than did plants collected from uninvaded areas. Accordingly, while there were fewer big squirreltail individuals in the invaded sites, they were more competitive with cheatgrass than were the more abundant conspecifics in nearby uninvaded areas. It is possible that annual grasses were the selective force that caused these population differences, which may contribute to the long-term persistence of the native populations. While it is tempting to restore degraded areas to higher densities of natives (usually done by bringing in outside seed material), such actions may impede long-term adaptation to new conditions by arresting or reversing the direction of ongoing natural selection in the resident population. If hot spots of rapid evolutionary change can be identified within invaded systems, these areas should be managed to promote desirable change and could serve as possible sources of restoration material or reveal traits that should be prioritized during the development of restoration seed material.

Genetic variation and local adaptation at a cheatgrass (Bromus tectorum) invasion edge in western Nevada

Cheatgrass (Bromus tectorum) is an invasive weed in western North America found primarily growing at elevations less than 2200 m. We asked whether cheatgrass is capable of becoming adapted to a marginal habitat, by investigating a population at a high elevation invasion edge. We used a combination of methods, including reciprocal field transplants, controlled environment studies and molecular analysis. High levels of SSR gene diversity (0.50 vs. 0.43) and comparable variation in phenotypic traits were observed at both the invasion edge and a low elevation, high‐density population. Three heterozygotes were observed in the edge population, which is unusual in this predominantly self‐pollinating plant. Plants from high elevations germinated more slowly in a growth chamber and had slower seedling growth rates. Survivorship was low at the edge (13%), compared with the low elevation site (55%), but surviving plants were of similar size and had equivalent reproductive output. Seed size positively affected survival and plant performance in the field and this trait was inherited. Emergence timing affected survival at the low elevation site and germination timing was also inherited. Local adaptation was seen in the low, rather than in the high elevation site, because of differential survival. While there was no evidence for local adaptation to the high elevation site observed in the field, family level and genotype‐level differences in traits that affected field performance, high genetic diversity at the invasion edge, and evidence of outcrossing in this highly selfing species indicates that the potential for adaptation to a marginal habitat exists within this population.

Competitive seedlings and inherited traits: a test of rapid evolution of Elymus multisetus (big squirreltail) in response to cheatgrass invasion.

Widespread invasion by Bromus tectorum (cheatgrass) in the Intermountain West has drastically altered native plant communities. We investigated whether Elymus multisetus (big squirreltail) is evolving in response to invasion and what traits contribute to increased performance. Seedlings from invaded areas exhibited significantly greater tolerance to B. tectorum competition and a greater ability to suppress B. tectorum biomass than seedlings from adjacent uninvaded areas. To identify potentially adaptive traits, we examined which phenological and phenotypic traits were correlated with seedling performance within the uninvaded area, determined their genetic variation by measuring sibling resemblance, and asked whether trait distribution had shifted in invaded areas. Increased tolerance to competition was correlated with early seedling root to shoot ratio, root fork number, and fine root length. Root forks differed among families, but none of these traits differed significantly across invasion status. Additionally, we surveyed more broadly for traits that varied between invaded and uninvaded areas. Elymus multisetus plants collected from invaded areas were smaller, allocated more biomass to roots, and produced a higher percentage of fine roots than plants from uninvaded areas. The ability of native populations to evolve in response to invasion has significant implications for the management and restoration of B. tectorum‐invaded communities.

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.

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.

Effects of experimental manipulation of light and nutrients on establishment of seedlings of native and invasive woody species in Long Island, NY forests

While earlier studies on the process of invasion often focused on single factors or on the general explanation of ‘disturbance,’ recent work has attempted to move towards a more mechanistic understanding of the factors that promote plant community invasion. Manipulative experiments provide a means for discerning causal relationships and interactive effects of environmental factors in promoting invasion; such experiments have been conducted in a number of grassland and shrub ecosystems. This study extends multifactor manipulative experiments into forest communities to compare factors influencing early seedling establishment for native and invasive woody plants. In Long Island, NY, invasion patterns are correlated with forest community type (pine barrens or hardwood), light availability, and soil N and Ca. We conducted manipulative field experiments in two different years to determine the relative importance and interaction of experimental gaps and N and Ca addition in pine barrens and hardwood forests in promoting invasion. We used seedlings of seven common native and invasive species in the first experiment, and 16 native and invasive species paired phylogenetically in the second experiment. Light had the strongest effect on plant growth; all plants grew more in gaps. We found no difference in the average growth rates of native and invasive species. Invasives responded more to high resources than did natives, with highest relative growth rates in gaps in the more fertile soils of the hardwood forests. Opportunities for invasion may differ from year to year, with differential success of invaders only in some years and under some environmental conditions. Clearly, to understand the complex interactions between resources and invasion in forests will require many manipulative experiments across a range of environments and using suites of invasive and native species.

Strong natural selection during plant restoration favors an unexpected suite of plant traits

Restoration is an opportunity to study natural selection: One can measure the distribution of traits in source propagules used to found populations, compare this with the distribution of traits in successful recruits, and determine the strength and direction of selection on potentially adaptive traits. We investigated whether natural selection influenced seedling establishment during postfire restoration in the Great Basin, an area where large‐scale restoration occurs with a few widely available cultivars planted over a large range of environmental conditions. We collected seeds from established plants of the perennial grass Elymus elymoides ssp. californicus (squirreltail) at two restoration sites and compared the distribution of phenotypic traits of surviving plants with the original pool of restoration seeds. Seeds were planted in common gardens for two generations. Plants grown from seeds that established in the field were a nonrandom subset of the original seeds, with directional selection consistently favoring a correlated suite of traits in both field sites: small plant and seed size, and earlier flowering phenology. These results demonstrate that natural selection can affect restoration establishment in strong and predictable ways and that adaptive traits in these sites were opposite of the current criteria used for selection of restoration material in this system.

The shifting balance of facilitation and competition affects the outcome of intra- and interspecific interactions over the life history of California grassland annuals

Trait-based resource competition in plants, wherein more similar plants compete more strongly for resources, is a foundation of niche-based explanations for the maintenance of diversity in plant communities. Alternatively, neutral theory predicts that community diversity can be maintained despite equivalent resource requirements among species. We examined interactions at three life history stages (germination, survival, and juvenile-adult growth) for three native and three exotic California annual species in a glasshouse experiment. We varied plant density and species composition in small pots, with pots planted with either intraspecific seeds or in a three species mix of intra- and interspecific neighbors. We saw a range of facilitative, neutral, and competitive interactions that varied significantly by species, rather than by native or exotic status. There were more competitive interactions at the emergence and juvenile-adult growth stages and more facilitative interactions for survival. Consequently, the relative strength of competition in intraspecific versus mixed-species communities depended on whether we considered only the juvenile-adult growth stage or the entire life history of the interacting plants. Using traditional analysis of juvenile-adult growth only, all species showed negative density-dependent interactions for final biomass production. However, when the net effect of plant interactions from seed to adult was considered, which is a prediction of population growth, two native species ceased to show negative density dependence, and the difference between intraspecific and mixed-species competition was only significant for one exotic species. Results were consistent with predictions of neutral, rather than niche, theory for five of six species.

Invasion Triangle: An Organizational Framework for Species Invasion

Species invasion is a complex, multifactor process. To encapsulate this complexity into an intuitively appealing, simple, and straightforward manner, we present an organizational framework in the form of an invasion triangle. The invasion triangle is an adaptation of the disease triangle used by plant pathologists to help envision and evaluate interactions among a host, a pathogen, and an environment. Our modification of this framework for invasive species incorporates the major processes that result in invasion as the three sides of the triangle: (1) attributes of the potential invader; (2) biotic characteristics of a potentially invaded site; and (3) environmental conditions of the site. The invasion triangle also includes the impact of external influences on each side of the triangle, such as climate and land use change. This paper introduces the invasion triangle, discusses how accepted invasion hypotheses are integrated in this framework, describes how the invasion triangle can be used to focus research and management, and provides examples of application. The framework provided by the invasion triangle is easy to use by both researchers and managers and also applicable at any level of data intensity, from expert opinion to highly controlled experiments. The organizational framework provided by the invasion triangle is beneficial for understanding and predicting why species are invasive in specific environments, for identifying knowledge gaps, for facilitating communication, and for directing management in regard to invasive species.

What Seeds to Plant in the Great Basin? Comparing Traits Prioritized in Native Plant Cultivars and Releases with those that Promote Survival in the Field

ABSTRACT: Restoration in the Great Basin is typically a large-scale enterprise, with aerial, drill, and broadcast seeding of perennial species common after wildfires. Arid conditions and invasive plants are significant barriers to overcome, but relatively simple changes to seeds used for restoration may improve success. Here we summarize: 1) the composition of seed mixes used in recent postfire seedings in Nevada, 2) traits that were valued when cultivars and other native seed materials were named and released, and 3) traits that have been demonstrated to increase native perennial grass performance in invaded systems. A review of 420 seeding treatments on public shrublands in Nevada between 2006 and 2009 indicated that native perennial grasses and native shrubs were most frequently included in these projects, followed by exotic and native forbs, and lastly, exotic perennial grasses. Native perennial grasses made up the bulk of seeds used in these treatments, with multiple species of grasses (average of 3.4 species) typically seeded per treatment, while the richness of other functional groups in seed mixes was closer to 1 species per treatment. Traits prioritized in cultivars and native seed material releases included, in order of frequency: forage quality and yield, seed yield, seedling vigor, ability to establish and persist, and drought tolerance, with many other traits mentioned with less frequency. Traits that had consistent support for improving native perennial grass performance in the field were related to early phenology, small size, and higher root allocation. Further tests to determine which traits improve shrub and forb establishment under field conditions could further refine seed source selection, and help maintain diversity in Great Basin systems.