Daniel Z. Atwater

Root contact responses and the positive relationship between intraspecific diversity and ecosystem productivity

We show that root-root recognition processes may help to explain how genetic diversity within species can lead to great ecosystem functioning. Roots of plants from the same population slowed down much more after contacting each other than roots of plants from different populations, suggesting that mixing genetic diversity may lead to greater root overlap and thereby greater ecosystem functioning.

An exotic invasive plant selects for increased competitive tolerance, but not competitive suppression, in a native grass

Exotic invasive plants can exert strong selective pressure for increased competitive ability in native plants. There are two fundamental components of competitive ability: suppression and tolerance, and the current paradigm that these components have equal influences on a species’ overall competitive ability has been recently questioned. If these components do not have equal influences on overall ability, then selection on competitive tolerance and suppression may be disproportionate. We used naturally invaded communities to study the effects of selection caused by an invasive forb, Centaurea stoebe, on a native grass, Pseudoroegneria spicata. P. spicata plants were harvested from within dense C. stoebe patches and from nearby uninvaded areas, divided clonally into replicates, then transplanted into a common garden where they grew alone or competed with C. stoebe. We found that P. spicata plants collected from within C. stoebe patches were significantly more tolerant of competition with C. stoebe than P. spicata plants collected from uninvaded areas, but plants from inside invaded patches were not superior at suppressing C. stoebe. These results are consistent with the hypothesis that strong competitors may select for tolerance to competition more than for the ability to suppress neighbors. This has important implications for how native plant communities may respond to invasion over time, and how invasive and native species may ultimately coexist.

Testing the mechanisms of diversity-dependent overyielding in a grass species.

Plant diversity enhances many ecosystem processes, including productivity, but these effects have been studied almost exclusively at the taxonomic scale of species. We explore the effect of intraspecific diversity on the productivity of a widespread and dominant grassland species using accessions collected from populations throughout its range. We found that increasing population/ecotype diversity of Pseudoroegneria spicata increased productivity to a similar degree as that reported for species diversity. However, we did not find evidence that overyielding was related to either resource depletion or to pathogenic soil fungi, two causes of overyielding in species-diverse communities. Instead, larger accessions overyielded at low diversity at the expense of smaller accessions, and small accessions overyielded through complementarity at all levels of diversity. Furthermore, overyielding was stronger for accessions from mesic environments, suggesting that local adaptation might predictably influence how plants respond to increases in diversity. This suggests that mass-based competition or other cryptic accession-specific processes had complex but important effects on overyielding. Our results indicate that the effects of diversity within a species can be substantial but that overyielding by intraspecifically diverse populations may not be through the same processes thought to cause overyielding in species diverse communities.

Multi-phase US spread and habitat switching of a post-columbian invasive, Sorghum halepense

Johnsongrass (Sorghum halepense) is a striking example of a post-Columbian founder event. This natural experiment within ecological time-scales provides a unique opportunity for understanding patterns of continent-wide genetic diversity following range expansion. Microsatellite markers were used for population genetic analyses including leaf-optimized Neighbor-Joining tree, pairwise FST, mismatch analysis, principle coordinate analysis, Tajima’s D, Fu’s F and Bayesian clusterings of population structure. Evidence indicates two geographically distant introductions of divergent genotypes, which spread across much of the US in <200 years. Based on geophylogeny, gene flow patterns can be inferred to have involved five phases. Centers of genetic diversity have shifted from two introduction sites separated by ~2000 miles toward the middle of the range, consistent with admixture between genotypes from the respective introductions. Genotyping provides evidence for a ‘habitat switch’ from agricultural to non-agricultural systems and may contribute to both Johnsongrass ubiquity and aggressiveness. Despite lower and more structured diversity at the invasion front, Johnsongrass continues to advance northward into cooler and drier habitats. Association genetic approaches may permit identification of alleles contributing to the habitat switch or other traits important to weed/invasive management and/or crop improvement.

Ecotypic diversity of a dominant grassland species resists exotic invasion

Many species are characterized by high levels of intraspecific or ecotypic diversity, yet we know little about how diversity within species influences ecosystem processes. Using a common garden experiment, we studied how intraspecific diversity within the widespread and often dominant North American native Pseudoroegneria spicata (Pursh) Á. Löve. affected invasion by Centaurea stoebe L. We experimentally manipulated Pseudoroegneria intraspecific diversity by changing the number of Pseudoroegneria ecotypes in common garden plots, using ecotypes collected throughout western North America. Invader biomass was 46% lower in mono-ecotype Pseudoroegneria plots than in control plots without any plants prior to invasion, and plots with 3–12 Pseudoroegneria ecotypes were 44% less invaded by Centaurea than the mono-ecotype plots. Across all plots, the total biomass of invading Centaurea plants was negatively correlated with total Pseudoroegneria biomass, but biotic resistance provided by high ecotypic diversity of Pseudoroegneria was not explained only by the increase in productivity that occurred with ecotypic diversity. Relative to Pseudoroegneria yield, Centaurea yield was lowest when Pseudoroegneria overyielded due to size-independent “complementarity” effects. This was not observed when overyielding was due to size-dependent effects. Our results suggest that the intraspecific diversity of a widespread and dominant species has the potential to impact invasion outcomes beyond its effects on native plant productivity and that mechanisms of biotic resistance to invaders may be to some degree independent of plant size.

Competition and propagule density affect sexual and clonal propagation of a weed

Many introduced species are capable of both sexual and vegetative reproduction. Our understanding of the ecology of such species depends on the trade-offs between vegetative and sexual reproduction and the ecological conditions that favor both modes of reproduction and how those factors influence the population ecology of introduced species. Here, we studied the efficacy of propagation via both seeds and rhizomes in Johnsongrass, a widespread invasive grass whose success is due to its prolific production of shattering seeds and rhizomes, the latter of which are readily dispersed by anthropogenic and natural processes. In a common garden in Virginia, we varied the density of seeds and rhizomes and manipulated whether recruits experienced interspecific competition. Johnsongrass recruited from both seeds and rhizomes. We compared the efficacy of seeds and rhizomes on a per propagule basis and by standardizing them according to their total carbon content. Rhizomes were more efficient than seeds on a per propagule basis, but seeds propagated more efficiently than rhizomes on a per unit of carbon basis, establishing in nearly all plots and obtaining much greater biomass than rhizomes. We also found that rhizomes were subject to stronger negative density dependence than seeds and were more sensitive to site variation and competition. Our results suggest that, provided sufficient dispersal, a single Johnsongrass plant produces enough propagules to establish over more than a hectare, even at relatively low propagule densities. Proper understanding of both seed and vegetative propagation is crucial for understanding the ecology of this and other invasive species that utilize multiple reproductive modes. Nomenclature: Johnsongrass; Sorghum halepense (L.) Pers. SORHA.

Evidence for fine-scale habitat specialisation in an invasive weed

As an exotic species colonises a new continent, it must overcome enormous environmental variation in its introduced range. Local adaptation of introduced species has frequently been observed at the continent scale, particularly in response to latitudinal climatic variation. However, significant environmental heterogeneity can also exist at the landscape scale. A small number of studies have provided evidence that introduced species may also be capable of phenotypic and genetic differentiation at much smaller spatial scales. For example, previously we found US agricultural and non-agricultural populations of Sorghum halepense (Johnsongrass) to be phenotypically and genetically distinct. In this study, we investigated whether this phenotypic differentiation of agricultural and non-agricultural populations of S. halepense is the result of fine-scale local specialisation. We surveyed a nationally collected S. halepense germplasm panel and also collected neighbouring agricultural and non-agricultural sub-populations of S. halepense at four sites throughout Western Virginia, USA, raising seedlings in common conditions mimicking both agricultural and non-agricultural habitats. At the national scale, we found evidence of habitat differentiation but not specialisation. However, at the local scale, we found evidence of specialisation in two of the four local populations to non-agricultural habitat, but no evidence of specialisation to agricultural habitat. These results show that local specialisation is a possible, but not guaranteed consequence of kilometre-scale habitat heterogeneity in invasive species. This finding contributes to a growing awareness of the importance of fine-scale local adaptation in the ecology and management of introduced and weedy species.