Comparisons of Efficiency of Two Formulations of Verbenone (4, 6, 6-trimethylbicyclo [3.1.1] hept-3-en-2-one) for Protecting Whitebark Pine, Pinus albicaulis (Pinales: Pinaceae) From Mountain Pine Beetle (Colopetera: Curculionidae).

Whitebark pine, Pinus albicaulis Engelm., is a subalpine tree endemic to western North America. This species provides multiple ecosystem services and is suffering widespread mortality from mountain pine beetle, Dendroctonus ponderosae Hopkins. Verbenone is a pheromone produced as D. ponderosae feed, and high air concentrations of verbenone deter D. ponderosae from colonizing trees. Synthetic verbenone has been formulated into products used to prevent D. ponderosae from colonizing trees. We compared the ability of verbenone pouches and SPLAT Verb to protect individuals and small stands of P. albicaulis. With individual trees in Montana, all treated trees survived regardless of verbenone formulation and rate, whereas untreated trees suffered 70 and 90% mortality in 2015 and 2016. In plot experiments in California from 2015 to 2017, and Oregon from 2015 to 2018, verbenone was applied to trees spaced ~10 m apart, and survival of small (12.7-23 cm DBH = diameter at 1.37 m height), medium (23.1-33 cm DBH) and large (>33 cm DBH) trees was compared. In California, where >80% of untreated trees survived, pouches increased survival ~2 to 3% and SPLAT Verb increased survival ~4 to 7% regardless of tree size. In Oregon, verbenone pouches and SPLAT Verb performed similarly on medium and small trees, but large trees had greater survival when treated with SPLAT Verb (~93%) than pouches (~82%). Compared to verbenone pouches, SPLAT Verb appears to better protect P. albicaulis from D. ponderosae.

Omitted variable bias in studies of plant interactions.

Models of plant-plant interactions underpin our understanding of species coexistence, invasive plant impacts, and plant community responses to climate change. In recent studies, models of competitive interactions failed predictive tests, thereby casting doubt on results of many past studies. We believe these model failures owe at least partly to heterogeneity in unmodeled factors (e.g., nutrients, soil pathogens) that affect both target plants and neighboring competitors. Such heterogeneity is ubiquitous, and models that do not account for it will suffer omitted variable bias. We used instrumental variables analysis to test for and correct omitted variable bias in studies that followed common protocols for measuring plant competition. In an observational study, omitted variables caused competition to seem like mutualism. In a quasi-experiment that partially controlled competitor abundances with seeding, omitted variables caused competition to seem about 35% weaker than it really was, even though the experiment occurred in an abandoned agricultural field where environmental heterogeneity was expected to be relatively low. Despite decades of research, consistently accurate estimates of competitive interactions remain elusive. The most foolproof way around this problem is true experiments that avoid omitted variable bias by completely controlling competitor abundances, but such experiments are rare.

Toward more robust plant-soil feedback research.

Understanding if and how plant-soil biota feedbacks (PSFs) shape plant communities has become a major research priority. In this paper, we draw on a recent, high-profile PSF study to illustrate that certain widely used experimental methods cannot reliably determine if PSFs occur. One problem involves gathering soil samples adjacent to multiple conditioning plants, mixing the samples and then growing phytometers in the mixtures to test for PSFs. This mixed soil approach does not establish that the conditioning plant being present caused the soil biota to be present, the first step of a PSF. Also, soil mixing approximates replacing raw data with averages prior to analysis, a move certain to generate falsely precise statistical estimates. False precision also results from sample sizes being artificially inflated when phytometers are misinterpreted as experimental units. Plant biomass ratios become another source of false precision when individual plant values contribute to multiple ratio observations. Any one of these common missteps can cause still living null hypotheses to be pronounced dead, and risks of this increase with numbers of missteps. If soil organisms truly structure plant communities, then null hypotheses indicating otherwise will not survive proper testing. We discuss conceptual, experimental and analytical refinements to facilitate accurate testing.

High precipitation and seeded species competition reduce seeded shrub establishment during dryland restoration.

Drylands comprise 40% of Earth's land mass and are critical to food security, carbon sequestration, and threatened and endangered wildlife. Exotic weed invasions, overgrazing, energy extraction, and other factors have degraded many drylands, and this has placed an increased emphasis on dryland restoration. The increased restoration focus has generated a wealth of experience, innovations and empirical data, yet the goal of restoring diverse, native, dryland plant assemblages composed of grasses, forbs, and shrubs has generally proven beyond reach. Of particular concern are shrubs, which often fail to establish or establish at trivially low densities. We used data from two Great Plains, USA coal mines to explore factors regulating shrub establishment. Our predictor data related to weather and restoration (e.g., seed rates, rock cover) variables, and our response data described shrub abundances on fields of the mines. We found that seeded non-shrubs, especially grasses, formed an important competitive barrier to shrub establishment: With every one standard deviation increase in non-shrub seed rate, the probability shrubs were present decreased ~0.1 and shrub cover decreased ~35%. Since new fields were seeded almost every year for > 20 years, the data also provided a unique opportunity to explore effects of stochastic drivers (i.e., precipitation, year effects). With every one standard deviation increase in precipitation the first growing season following seeding, the probability shrubs were present decreased ~0.07 and shrub cover decreased ~47%. High precipitation appeared to harm shrubs by increasing grass growth/competition. Also, weak evidence suggested shrub establishment was better in rockier fields where grass abundance/competition was lower. Multiple lines of evidence suggest reducing grass seed rates below levels typically used in Great Plains restoration would benefit shrubs without substantially impacting grass stand development over the long term. We used Bayesian statistics to estimate effects of seed rates and other restoration predictors probabilistically to allow knowledge of the predictors' effects to be refined through time in an adaptive management framework. We believe this framework could improve restoration planning in a variety of systems where restoration outcomes remain highly uncertain and ongoing restoration efforts are continually providing new data of value for reducing the uncertainty.

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.

Predicting plant responses to mycorrhizae: integrating evolutionary history and plant traits.

We assessed whether (1) arbuscular mycorrhizal colonization of roots (RC) and/or plant responses to arbuscular mycorrhizae (MR) vary with plant phylogeny and (2) MR and RC can be more accurately predicted with a phylogenetic predictor relative to a null model and models with plant trait and taxonomic predictors. In a previous study, MR and RC of 95 grassland species were measured. We constructed a phylogeny for these species and found it explained variation in MR and RC. Next, we used multiple regressions to identify the models that most accurately predicted plant MR. Models including either phylogenetic or phenotypic and taxonomic information similarly improved our ability to predict MR relative to a null model. Our study illustrates the complex evolutionary associations among species and constraints of using phylogenetic information, relative to plant traits, to predict how a plant species will interact with AMF.

Control effort exacerbates invasive-species problem.

Ecosystem managers face a difficult decision when managing invasive species. If they use aggressive practices to reduce invader abundances, they will likely reduce invaders' competitive impacts on natives. But it is often difficult or impossible to reduce invaders without damaging natives. So a critical question becomes: Which is worse for native biota, invaders or things done to control invaders? We attempted to answer this question for a common scenario. We studied several grassland natives exhibiting long-term coexistence with an invader and asked how aggressive management (herbicide use) affected the natives. Whether or not grazing was excluded, one-time herbicide use made two native forbs exceedingly rare for our entire 16-year study period. Herbicide also made several other native forbs rare, but only when grazing was excluded, and there is evidence that the dominant invader became more abundant in response to the decreases in native-forb abundances. Throughout the world, terrestrial and aquatic ecosystems are receiving herbicide applications for exotic-species control. Some of the applications are doubtless warranted because they target small invader patches or larger areas with virtually no remaining natives. However, other herbicide applications occur where large native populations occur, and our data suggest that these applications can be ill advised. Our cautionary tale is told using an herbicide-treated grassland, but our results should be considered wherever invasive-species management damages native species.

Grassland invader responses to realistic changes in native species richness.

The importance of species richness for repelling exotic plant invasions varies from ecosystem to ecosystem. Thus, in order to prioritize conservation objectives, it is critical to identify those ecosystems where decreasing richness will most greatly magnify invasion risks. Our goal was to determine if invasion risks greatly increase in response to common reductions in grassland species richness. We imposed treatments that mimic management-induced reductions in grassland species richness (i.e., removal of shallow- and/or deep-rooted forbs and/or grasses and/or cryptogam layers). Then we introduced and monitored the performance of a notorious invasive species (i.e., Centaurea maculosa). We found that, on a per-gram-of-biomass basis, each resident plant group similarly suppressed invader growth. Hence, with respect to preventing C. maculosa invasions, maintaining overall productivity is probably more important than maintaining the productivity of particular plant groups or species. But at the sites we studied, all plant groups may be needed to maintain overall productivity because removing forbs decreased overall productivity in two of three years. Alternatively, removing forbs increased productivity in another year, and this led us to posit that removing forbs may inflate the temporal productivity variance as opposed to greatly affecting time-averaged productivity. In either case, overall productivity responses to single plant group removals were inconsistent and fairly modest, and only when all plant groups were removed did C. maculosa growth increase substantially over a no-removal treatment. As such, it seems that intense disturbances (e.g., prolonged drought, overgrazing) that deplete multiple plant groups may often be a prerequisite for C. maculosa invasion.