Yvette K. Ortega

WEAK VS. STRONG INVADERS OF NATURAL PLANT COMMUNITIES: ASSESSING INVASIBILITY AND IMPACT

In response to the profound threat of exotic species to natural systems, much attention has been focused on the biotic resistance hypothesis, which predicts that diverse communities should better resist invasions. While studies of natural communities generally refute this hypothesis, reporting positive relationships between native species diversity and invasibility, some local-scale studies have instead obtained negative relationships. Most treatments of the topic have failed to recognize that all exotic invaders do not behave alike: while “weak” invaders become minor components of communities, “strong” invaders become community dominants at the expense of native species. At the same time, the specific impacts of strong invaders on communities are poorly documented yet critical to understanding implications of diversity loss. With these shortfalls in mind, we examined local-scale relationships between native and exotic plant taxa in bunchgrass communities of western Montana, USA. We found that measures of...

Invasion of an exotic forb impacts reproductive success and site fidelity of a migratory songbird.

Although exotic plant invasions threaten natural systems worldwide, we know little about the specific ecological impacts of invaders, including the magnitude of effects and underlying mechanisms. Exotic plants are likely to impact higher trophic levels when they overrun native plant communities, affecting habitat quality for breeding songbirds by altering food availability and/or nest predation levels. We studied chipping sparrows (Spizella passerina) breeding in savannas that were either dominated by native vegetation or invaded by spotted knapweed (Centaurea maculosa), an exotic forb that substantially reduces diversity and abundance of native herbaceous plant species. Chipping sparrows primarily nest in trees but forage on the ground, consuming seeds and arthropods. We found that predation rates did not differ between nests at knapweed and native sites. However, initiation of first nests was delayed at knapweed versus native sites, an effect frequently associated with low food availability. Our seasonal fecundity model indicated that breeding delays could translate to diminished fecundity, including dramatic declines in the incidence of double brooding. Site fidelity of breeding adults was also substantially reduced in knapweed compared to native habitats, as measured by return rates and shifts in territory locations between years. Declines in reproductive success and site fidelity were greater for yearling versus older birds, and knapweed invasion appeared to exacerbate differences between age classes. In addition, grasshoppers, which represent an important prey resource, were substantially reduced in knapweed versus native habitats. Our results strongly suggest that knapweed invasion can impact chipping sparrow populations by reducing food availability. Food chain effects may be an important mechanism by which strong plant invaders impact songbirds and other consumers.

Fire and mice: seed predation moderates fire's influence on conifer recruitment.

In fire-adapted ecosystems, fire is presumed to be the dominant ecological force, and little is known about how consumer interactions influence forest regeneration. Here, we investigated seed predation by deer mice (Peromyscus maniculatus) and its effects on recruitment of ponderosa pine (Pinus ponderosa) and Douglas-fir (Pseudotsuga menziesii) seedlings in unburned and recently burned fire-adapted montane forests in west-central Montana, USA. Deer mice were almost twice as abundant in burned than unburned stands. Deer mouse removal of seeds from petri dishes was two times higher in burned than in unburned stands, and seed removal levels were 8% higher for ponderosa pine than for the smaller Douglas-fir seeds. In seed-addition experiments, emergence of seedlings in deer mouse-exclusion cages was almost six times higher in burned compared to unburned forest. In both burned and unburned forest, emergence was lower for ponderosa pine than for Douglas-fir. Seedling survival to establishment did not differ between conifer species but was considerably higher in burned than in unburned forest. However, effects of seed predation on recruitment prevailed over fire effects: in cages allowing access by deer mice, emergence and establishment were extremely rare for both conifer species in both burned and unburned forest. This research suggests that consumer interactions can substantially influence recruitment even in fire-adapted forest ecosystems.

Effects of biological control agents and exotic plant invasion on deer mouse populations

Exotic insects are commonly introduced as biological control agents to reduce densities of invasive exotic plants. Although current biocontrol programs for weeds take precautions to minimize ecological risks, little attention is paid to the potential nontarget effects of introduced food subsidies on native consumers. Previous research demonstrated that two gall flies (Urophora affinis and U. quadrifasciata) introduced for biological control of spotted knapweed (Centaurea maculosa) dramatically affect the foraging ecology of the deer mouse (Peromyscus maniculatus), a native generalist predator with important trophic linkages. In the current study, we found that relative abundance of deer mice was elevated twofold in grassland habitats with high densities of spotted knapweed and gall fly food sources, compared to those dominated by native vegetation, in two of three years. Availability of gall fly larvae during the critical overwinter period appeared to reduce overwinter population declines of mice in knapweed-invaded habitats. These positive effects on populations apparently overshadowed negative effects on breeding productivity associated with knapweed invasion and loss of the gall fly resource during the summer. Our results suggest that insect biocontrol agents can subsidize native consumer populations, setting the stage for various indirect effects on food webs. Comprehensive understandings of the conditions under which introduced biological control agents may exhibit nontarget effects on native food webs are needed to further develop criteria for screening potential biocontrol agents before they are released.

Long-Term Effects of Weed Control With Picloram Along a Gradient of Spotted Knapweed Invasion

Abstract Broadleaf herbicides are commonly used in rangelands to suppress exotic weeds and release native communities from negative impacts of invasion. However, few studies have comprehensively evaluated treatment effects on differing community components across a gradient of initial invasion levels. We conducted a 6-yr experiment within grasslands of western Montana to measure local-scale effects of a broadcast application of picloram on 1) cover of the target invader, spotted knapweed (Centaurea stoebe L.), 2) prevalence of native functional groups, and 3) the secondary invader cheatgrass (Bromus tectorum L.) at differing initial levels of knapweed invasion. Treatment effectively suppressed knapweed, with cover in treated vs. control plots reduced by > 60% in the sixth posttreatment year. Treatment also appeared to alleviate knapweeds impacts on native perennial grasses, but only at the highest initial level of invasion, where cover of this group increased by > 30% in treated vs. control plots to equal levels associated with noninvaded plots. In some cases, treatment appeared to exacerbate knapweeds impacts on native forbs. At the no-invasion level, perennial forb cover declined by > 20% in treated vs. control plots to match values associated with moderate or high levels of invasion, but these treatment effects were minimal at the latter invasion levels. Across initial invasion levels, species richness of perennial and/or annual forbs declined by > 20% in treated vs. control plots. Treatment also promoted increases in cheatgrass cover, although differences between treated and control plots were relatively small by the sixth posttreatment year. Overall, effects of picloram application depended on initial levels of knapweed invasion, largely due to the varying strength of release effects. Selective treatment of invaded patches vs. broadcast applications would reduce side effects of broadleaf herbicide application and increase compatibility with other management measures designed to improve rangeland conditions and restore grassland communities.

Darwin’s naturalization hypothesis up-close: Intermountain grassland invaders differ morphologically and phenologically from native community dominants

Darwin’s naturalization hypothesis predicts that successful invaders will tend to differ taxonomically from native species in recipient communities because less related species exhibit lower niche overlap and experience reduced biotic resistance. This hypothesis has garnered substantial support at coarse scales. However, at finer scales, the influence of traits and niche use on invasibility and invader impacts is poorly understood. Within grasslands of western Montana, USA, we compared morphological and phenological traits for five top exotic invasive forbs and five dominant native forbs using multivariate techniques to examine niche separation between exotics and natives. Exotic forbs differed from native forbs in multivariate space. Phenologically, native forbs synchronized vegetative growth with bolting and flowering early in spring. In contrast, exotics initiated vegetative growth concurrent with natives but bolted and flowered later. Morphologically, vegetative growth of exotics was three times shorter and narrower, but flowering stem growth was 35% taller and 65% wider than the natives. Collectively, these patterns suggest different strategies of resource uptake and allocation. Additionally, following wildfire, survival was four times higher for exotics compared to natives, and three times more of the surviving exotics flowered. The exotics we examined appeared to be exploiting an empty community-level niche. The resulting pattern of trait differences between exotics and natives suggests a predictable pattern of invasion and a predictable trajectory of community change. Our results illustrate how quantifying trait differences between invading exotics and natives at the within-community scale can improve understandings of community invasibility and invader impacts.

Effects of Picloram Application on Community Dominants Vary With Initial Levels of Spotted Knapweed (Centaurea stoebe) Invasion

Abstract Broadleaf herbicides are commonly used to suppress exotic weeds with the intent of releasing native species from negative impacts of invasion. However, weed control measures can also have unintended consequences that should be considered along with intended effects. We conducted a controlled field experiment within bunchgrass communities of western Montana to examine if broadcast application of the broadleaf herbicide, picloram, may mitigate impacts of the exotic forb, spotted knapweed, on the dominant native grass, bluebunch wheatgrass, and forb, arrowleaf balsamroot. Local-scale relationships between native species and spotted knapweed cover served as a baseline for evaluating treatment effects at differing spotted knapweed invasion levels. To examine secondary invasion, we also measured treatment effects on the exotic grass, downy brome, relative to initial levels of spotted knapweed cover. Picloram application suppressed spotted knapweed cover by 70 to 80%. Treatment appeared to release cover and seed production of bluebunch wheatgrass, causing increases that varied positively with initial spotted knapweed cover. Bluebunch wheatgrass measures were elevated by as much as fourfold in treated vs. control plots, exceeding baseline levels in noninvaded plots. For arrowleaf balsamroot, negative effects of treatment prevailed, particularly where initial spotted knapweed cover was low. Arrowleaf balsamroot cover and fecundity variables were reduced by as much as 60% in treated vs. control plots, to levels typifying baseline conditions in highly invaded plots. In addition, treatment released downy brome, with cover increases from 2- to 20-fold. A controlled experiment selectively removing spotted knapweed showed similar release of downy brome. Our results show that picloram effects can depend on initial levels of weed invasion and may include substantial side effects, particularly when broadcast applications are used. Integrated approaches that include seeding of desirable species may be needed to enhance plant community resistance to secondary invaders and reinvasion by the target weed. Nomenclature: Picloram; downy brome, Bromus tectorum L. BRTE; spotted knapweed, Centaurea stoebe L. CEST8; arrowleaf balsamroot, Balsamorhiza sagittata (Pursh) Nutt. BASA3; bluebunch wheatgrass, Pseudoroegneria spicata (Pursh) A. Löve PSSP6.

Quantifying “apparent” impact and distinguishing impact from invasiveness in multispecies plant invasions

The quantification of invader impacts remains a major hurdle to understanding and managing invasions. Here, we demonstrate a method for quantifying the community-level impact of multiple plant invaders by applying Parker et al.s (1999) equation (impact = range x local abundance x per capita effect or per unit effect) using data from 620 survey plots from 31 grasslands across west-central Montana, USA. In testing for interactive effects of multiple invaders on native plant abundance (percent cover), we found no evidence for invasional meltdown or synergistic interactions for the 25 exotics tested. While much concern exists regarding impact thresholds, we also found little evidence for nonlinear relationships between invader abundance and impacts. These results suggest that management actions that reduce invader abundance should reduce invader impacts monotonically in this system. Eleven of 25 invaders had significant per unit impacts (negative local-scale relationships between invader and native cover). In decomposing the components of impact, we found that local invader abundance had a significant influence on the likelihood of impact, but range (number of plots occupied) did not. This analysis helped to differentiate measures of invasiveness (local abundance and range) from impact to distinguish high-impact invaders from invaders that exhibit negligible impacts, even when widespread. Distinguishing between high- and low-impact invaders should help refine trait-based prediction of problem species. Despite the unique information derived from evaluation of per unit effects of invaders, invasiveness scores based on range and local abundance produced similar rankings to impact scores that incorporated estimates of per unit effects. Hence, information on range and local abundance alone was sufficient to identify problematic plant invaders at the regional scale. In comparing empirical data on invader impacts to the state noxious weed list, we found that the noxious weed list captured 45% of the high impact invaders but missed 55% and assigned the lowest risk category to the highest-impact invader. While such subjective weed lists help to guide invasive species management, empirical data are needed to develop more comprehensive rankings of ecological impacts. Using weed lists to classify invaders for testing invasion theory is not well supported.

Negative plant‐soil feedbacks increase with plant abundance, and are unchanged by competition

Plant-soil feedbacks and interspecific competition are ubiquitous interactions that strongly influence the performance of plants. Yet few studies have examined whether the strength of these interactions corresponds with the abundance of plant species in the field, or whether feedbacks and competition interact in ways that either ameliorate or exacerbate their effects in isolation. We sampled soil from two intermountain grassland communities where we also measured the relative abundance of plant species. In greenhouse experiments, we quantified the direction and magnitude of plant-soil feedbacks for 10 target species that spanned a range of abundances in the field. In soil from both sites, plant-soil feedbacks were mostly negative, with more abundant species suffering greater negative feedbacks than rare species. In contrast, the average response to competition for each species was unrelated with its abundance in the field. We also determined how competitive response varied among our target species when plants competed in live vs. sterile soil. Interspecific competition reduced plant size, but the strength of this negative effect was unchanged by plant-soil feedbacks. Finally, when plants competed interspecifically, we asked how conspecific-trained, heterospecific-trained, and sterile soil influenced the competitive responses of our target species and how this varied depending on whether target species were abundant or rare in the field. Here, we found that both abundant and rare species were not as harmed by competition when they grew in heterospecific-trained soil compared to when they grew in conspecific-cultured soil. Abundant species were also not as harmed by competition when growing in sterile vs. conspecific-trained soil, but this was not the case for rare species. Our results suggest that abundant plants accrue species-specific soil pathogens to a greater extent than rare species. Thus, negative feedbacks may be critical for preventing abundant species from becoming even more abundant than rare species.

Trap-induced mass declines in small mammals: Mass as a population index

Body mass is routinely used as an index of physical condition for comparing small-mammal populations. However, trapping effects on animals may undermine the effectiveness of body mass as an index of population health. We examined the effects of live-trapping on body mass of 3 small-mammal species: deer mice (Peromyscus maniculatus), southern red-backed voles (Clethrionomys gapperi), and red-tailed chipmunks (Tamias ruficaudus). We found that live-trapping resulted in significant trap-induced body mass declines (TMDs) that varied by species, age, and sex. Longer rest intervals between captures reduced TMDs in deer mice, but not in other species. The TMDs were more dramatic for deer mice that died in traps than for those that did not, suggesting that TMDs may induce trap mortality and reduce survivorship in small mammals that are live-trapped. The proximate causes of TMDs remain unknown, but dehydration and hypothermia are likely causal agents. Since TMDs vary by species, age, and sex, and exposure to heating and cooling vary in space and time (e.g., by habitat and season), body mass as measured by live-trapping appears to be a negatively biased index whose deviation from true mass may vary among study populations and demographic groups. Researchers can attempt to bias correct indices of body mass using regression approaches, but such adjustments require large samples to be robust and so may not necessarily improve the raw indices. Additional research is needed to identify the causes of TMDs to better understand biases associated with using body mass as a population index and minimize impacts on small mammals.