Diane L. Larson

ALIEN PLANT INVASION IN MIXED-GRASS PRAIRIE: EFFECTS OF VEGETATION TYPE AND ANTHROPOGENIC DISTURBANCE

The ability of alien plant species to invade a region depends not only on attributes of the plant, but on characteristics of the habitat being invaded. Here, we examine characteristics that may influence the success of alien plant invasion in mixed-grass prairie at Theodore Roosevelt National Park, in western North Dakota, USA. The park consists of two geographically separate units with similar vegetation types and management history, which allowed us to examine the effects of native vegetation type, anthropogenic distur- bance, and the separate park units on the invasion of native plant communities by alien plant species common to counties surrounding both park units. If matters of chance related to availability of propagules and transient establishment opportunities determine the success of invasion, park unit and anthropogenic disturbance should better explain the variation in alien plant frequency. If invasibility is more strongly related to biotic or physicaL char- acteristics of the native plant communities, models of alien plant occurrence should include vegetation type as an explanatory variable. We examined >1300 transects across all veg- etation types in both units of the park. Akaikes Information Criterion (AIC) indicated that the fully parameterized model, including the interaction among vegetation type, disturbance, and park unit, best described the distribution of both total number of alien plants per transect and frequency of alien plants on transects where they occurred. Although all vegetation types were invaded by alien plants, mesic communities had both greater numbers and higher frequencies of alien plants than did drier communities. A strong element of stochasticity, reflected in differences in frequencies of individual species between the two park units, suggests that prediction of risk of invasion will always involve uncertainty. In addition, despite well-documented associations between anthropogenic disturbance and alien plant invasion, five of the six most abundant alien species at Theodore Roosevelt National Park had distributions unrelated to disturbance. We recommend that vegetation type be explicitly taken into account when designing monitoring plans for alien species in natural areas.

Soil modification by invasive plants: effects on native and invasive species of mixed-grass prairies

Invasive plants are capable of modifying attributes of soil to facilitate further invasion by conspecifics and other invasive species. We assessed this capability in three important plant invaders of grasslands in the Great Plains region of North America: leafy spurge (Euphorbia esula), smooth brome (Bromus inermis) and crested wheatgrass (Agropyron cristatum). In a glasshouse, these three invasives or a group of native species were grown separately through three cycles of growth and soil conditioning in both steam-pasteurized and non-pasteurized soils, after which we assessed seedling growth in these soils. Two of the three invasive species, Bromus and Agropyron, exhibited significant self-facilitation via soil modification. Bromus and Agropyron also had significant facilitative effects on other invasives via soil modification, while Euphorbia had significant antagonistic effects on the other invasives. Both Agropyron and Euphorbia consistently suppressed growth of two of three native forbs, while three native grasses were generally less affected. Almost all intra- and interspecific effects of invasive soil conditioning were dependent upon presence of soil biota from field sites where these species were successful invaders. Overall, these results suggest that that invasive modification of soil microbiota can facilitate plant invasion directly or via ‘cross-facilitation’ of other invasive species, and moreover has potential to impede restoration of native communities after removal of an invasive species. However, certain native species that are relatively insensitive to altered soil biota (as we observed in the case of the forb Linum lewisii and the native grasses), may be valuable as ‘nurse’species in restoration efforts.

Effects of the herbicide atrazine on Ambystoma tigrinum metamorphosis: duration, larval growth, and hormonal response

We exposed larval tiger salamanders (Ambystoma tigrinum) reared in the laboratory from eggs collected from a prairie wetland in North Dakota to three concentrations of atrazine (0, 75, and 250 mg/L) in a static renewal test to determine the pesticides effect on (1) plasma corticosterone and thyroxine concentrations, (2) larval size, and (3) days‐to‐stage at stages 2 and 4 of metamorphic climax. We found significant effects of atrazine on each of these response variables. Plasma thyrox‐ine was elevated in both atrazine‐exposed groups compared with the control group; plasma corticosterone was depressed in the 75 mg/L treatment compared with both the control and 250 mg/L treatment. Larvae exposed to 75 mg/L atrazine reached stage 4 later but at a size and weight comparable to the control group. By contrast, larvae in the 250 mg/L treatment progressed to stage 4 at the same time but at a smaller size and lower weight than larvae in the control group. These results indicate that the herbicide has the potential to influence tiger salamander life history. We present a model consistent with our results, whereby corticosterone and thyroxine interact to regulate metamorphosis of tiger salamanders based on nutrient assimilation and adult fitness.

Native weeds and exotic plants: relationships to disturbance in mixed-grass prairie*

Disturbance frequently is implicated in the spread of invasive exotic plants. Disturbances may be broadly categorized as endogenous (e.g., digging by fossorial animals) or exogenous (e.g., construction and maintenance of roads and trails), just as weedy species may be native or exotic in origin. The objective of this study was to characterize and compare exotic and native weedy plant occurrence in and near three classes of disturbance – digging by prairie dogs (an endogenous disturbance to which native plants have had the opportunity to adapt), paved or gravel roads (an exogenous disturbance without natural precedent), and constructed trails (an exogenous disturbance with a natural precedent in trails created by movement of large mammals) – in three geographically separate national park units. I used plant survey data from the North and South Units of Theodore Roosevelt National Park and Wind Cave National Park in the northern mixed-grass prairie of western North and South Dakota, USA, to characterize the distribution of weedy native and exotic plants with respect to the three disturbance classes as well as areas adjacent to them. There were differences both in the susceptibility of the disturbance classes to invasion and in the distributions of native weeds and exotic species among the disturbance classes. Both exotic and native weedy species richness were greatest in prairie dog towns and community composition there differed most from undisturbed areas. Exotic species were more likely to thrive near roadways, where native weedy species were infrequently encountered. Exotic species were more likely to have spread beyond the disturbed areas into native prairie than were weedy native species. The response of individual exotic plant species to the three types of disturbance was less consistent than that of native weedy species across the three park units.

Temporal dynamics of leafy spurge (Euphorbia esula) and two species of flea beetles (Aphthona spp.) used as biological control agents

Abstract The goal of this study was to evaluate the biological control program of leafy spurge ( Euphorbia esula ) in a large natural area, Theodore Roosevelt National Park, western North Dakota, USA. Aphthona lacertosa and Aphthona nigriscutis have been released at more than 1800 points in the 18,600-ha South Unit of the park beginning in 1989; most releases have occurred since 1994. We established permanent vegetation plots throughout the infested area of the park and determined stem counts and biomass of leafy spurge and abundance of the two flea beetle species at these plots each year from 1999 to 2001. Both biomass and stem counts declined over the 3 years of the study. Both species of flea beetle are well established within the park and have expanded into areas where they were not released. A. nigriscutis was more abundant than A. lacertosa in the grassland areas we surveyed, but in all other habitats abundances were similar. Using structural equation models, only A. lacertosa could be shown to have a significant effect on counts of mature stems of leafy spurge. A. nigriscutis numbers were positively correlated with stem counts of mature stems. Previous year’s stem counts had the greatest influence on change in stem counts over each 2-year time step examined with structural equation models.

Soil–Occupancy Effects of Invasive and Native Grassland Plant Species on Composition and Diversity of Mycorrhizal Associations

Abstract Diversified grasslands that contain native plant species can produce biofuels, support sustainable grazing systems, and produce other ecosystem services. However, ecosystem service production can be disrupted by invasion of exotic perennial plants, and these plants can have soil-microbial “legacies” that may interfere with establishment and maintenance of diversified grasslands even after effective management of the invasive species. The nature of such legacies is not well understood, but may involve suppression of mutualisms between native species and soil microbes. In this study, we tested the hypotheses that legacy effects of invasive species change colonization rates, diversity, and composition of arbuscular-mycorrhizal fungi (AMF) associated with seedlings of co-occurring invasive and native grassland species. In a glasshouse, experimental soils were conditioned by cultivating three invasive grassland perennials, three native grassland perennials, and a native perennial mixture. Each was grown separately through three cycles of growth, after which we used T-RFLP analysis to characterize AMF associations of seedlings of six native perennial and six invasive perennial species grown in these soils. Legacy effects of soil conditioning by invasive species did not affect AMF richness in seedling roots, but did affect AMF colonization rates and the taxonomic composition of mycorrhizal associations in seedling roots. Moreover, native species were more heavily colonized by AMF and roots of native species had greater AMF richness (number of AMF operational taxonomic units per seedling) than did invasive species. The invasive species used to condition soil in this experiment have been shown to have legacy effects on biomass of native seedlings, reducing their growth in this and a previous similar experiment. Therefore, our results suggest that successful plant invaders can have legacies that affect soil-microbial associations of native plants and that these effects can inhibit growth of native plant species in invaded communities. Management Implications: In the midwestern US, regulatory, market and policy pressures could convert large areas from annual agriculture to semi-natural grassland agroecosystems, e.g., as part of a national effort to produce energy crops. Native grassland perennials could be used in these grasslands to reduce production costs, conserve soil quality, conserve native biodiversity, and enhance carbon sequestration in grassland agroecosystems. However, producer interest in semi-natural grassland systems is reduced by current difficulties in reliable and cost-effective establishment of these species, and weed management during establishment is a major concern. Many lines of evidence suggest that weedy exotic species can alter soils physically and/or microbially, creating a “legacy” that persists after control or removal of these species. This legacy effect may contribute significantly to the risk of additional weed invasion and poor performance of desirable species during grassland establishment. In smooth brome, crested wheatgrass and leafy spurge, three exotic perennials that are highly invasive in grasslands, we examined legacy effects on AMF associations of a set of native and invasive species in these grasslands. Smooth brome, crested wheatgrass and leafy spurge had legacy effects that altered AMF colonization rates and community composition in seedling roots of native and invasive species, In a previous analysis of these data (Jordan et al. 2011), we found that these invasive species also had legacy effects that reduced growth of some native species. However, these legacy effects on growth and AMF colonization of natives varied substantially among these invasive species, suggesting that such effects are not uniform. Taken together, these legacy effects on AMF associations and growth suggest that restoration of AMF communities in soil may be important to cost-effective establishment and weed management in semi-natural grassland agroecosystems.

Role of Invasive Melilotus officinalis in Two Native Plant Communities

This study examines the impact of the exotic nitrogen-fixing legume Melilotus officinalis (L.) Lam. on native and exotic species cover in two Great Plains ecosystems in Badlands National Park, South Dakota. Melilotus is still widely planted and its effects on native ecosystems are not well studied. Melilotus could have direct effects on native plants, such as through competition or facilitation. Alternatively, Melilotus may have indirect effects on natives, e.g., by favoring exotic species which in turn have a negative effect on native species. This study examined these interactions across a 4-year period in two contrasting vegetation types: Badlands sparse vegetation and western wheatgrass (Pascopyrum smithii) mixed-grass prairie. Structural equation models were used to analyze the pathways through which Melilotus, native species, and other exotic species interact over a series of 2-year time steps. Melilotus can affect native and exotic species both in the current year and in the years after its death (a lag effect). A lag effect is possible because the death of a Melilotus plant can leave an open, potentially nitrogen-enriched site on the landscape. The results showed that the relationship between Melilotus and native and exotic species varied depending on the habitat and the year. In Badlands sparse vegetation, there was a consistent, strong, and positive relationship between Melilotus cover and native and exotic species cover suggesting that Melilotus is acting as a nurse plant and facilitating the growth of other species. In contrast, in western wheatgrass prairie, Melilotus was acting as a weak competitor and had no consistent effect on other species. In both habitats, there was little evidence for a direct lag effect of Melilotus on other species. Together, these results suggest both facilitative and competitive roles for Melilotus, depending on the vegetation type it invades.

Evidence of Qualitative Differences between Soil-Occupancy Effects of Invasive vs. Native Grassland Plant Species

Abstract Diversified grasslands that contain native plant species are being recognized as important elements of agricultural landscapes and for production of biofuel feedstocks as well as a variety of other ecosystem services. Unfortunately, establishment of such grasslands is often difficult, unpredictable, and highly vulnerable to interference and invasion by weeds. Evidence suggests that soil-microbial “legacies” of invasive perennial species can inhibit growth of native grassland species. However, previous assessments of legacy effects of soil occupancy by invasive species that invade grasslands have focused on single invasive species and on responses to invasive soil occupancy in only a few species. In this study, we tested the hypothesis that legacy effects of invasive species differ qualitatively from those of native grassland species. In a glasshouse, three invasive and three native grassland perennials and a native perennial mixture were grown separately through three cycles of growth and soil conditioning in soils with and without arbuscular mycorrhizal fungi (AMF), after which we assessed seedling growth in these soils. Native species differed categorically from invasives in their response to soil conditioning by native or invasive species, but these differences depended on the presence of AMF. When AMF were present, native species largely had facilitative effects on invasive species, relative to effects of invasives on other invasives. Invasive species did not facilitate native growth; neutral effects were predominant, but strong soil-mediated inhibitory effects on certain native species occurred. Our results support the hypothesis that successful plant invaders create biological legacies in soil that inhibit native growth, but suggest also this mechanism of invasion will have nuanced effects on community dynamics, as some natives may be unaffected by such legacies. Such native species may be valuable as nurse plants that provide cost-effective restoration of soil conditions needed for efficient establishment of diversified grasslands. Interpretive Summary: In the Midwestern United States, regulatory, market, and policy pressures could convert large areas from annual agriculture to seminatural grassland agroecosystems, e.g., as part of a national effort to produce energy crops. Native grassland perennials could be used in these grasslands to reduce production costs, conserve soil quality, conserve native biodiversity, and enhance carbon sequestration in grassland agroecosystems. However, producer interest in seminatural grassland systems is reduced by current difficulties in reliable and cost-effective establishment of these species, and weed management during establishment is a major concern. Many lines of evidence suggest that weedy exotic species can alter soils physically, microbially, or both, creating a “legacy: that persists after control or removal of these species. This legacy effect may contribute significantly to the risk of additional weed invasion and poor performance of desirable species during grassland establishment. We examined such legacy effects of smooth brome, crested wheatgrass, and leafy spurge, three exotic perennials that are highly invasive in grasslands. These species had strong inhibitory legacy effects on certain native species, but other native species were unaffected. We found that native species did not have inhibitory legacy effects, suggesting that managers should expect that successful plant invaders may leave soil legacies that will inhibit native growth in the establishment phase. We found that some natives were unaffected by such legacies, and these particular species may be valuable as “nurse plants” or cover crops that provide cost-effective conditioning of soils, thereby restoring soil conditions needed for efficient establishment of desirable native species.

Canada Thistle Biological Control Agents on Two South Dakota Wildlife Refuges

Abstract We monitored populations of Canada thistle biocontrol agents Cassida rubiginosa, Ceutorhynchus litura, Larinus (= Hadroplantus) planus, Urophora cardui, Orellia (= Terellia) ruficauda, and Rhinocyllus conicus on Canada thistle (Cirsium arvense) at two national wildlife refuges in South Dakota from 1999 through 2003. C. litura, U. cardui, O. ruficauda, and R. conicus were present on both refuges. Agent populations were low except for C. litura, which was present in up to 90% of stems in some plots. C. litura infestation did not reduce thistle flowering, stem length, or over-winter survival. There was no change in thistle stem numbers over the study period and no difference in stem numbers in areas of high C. litura populations compared to areas of low C. litura populations. Our results suggest that insect biological control agents are inadequate for reduction of Canada thistle in southern South Dakota.

Potential Effects of Anthropogenic Greenhouse Gases on Avian Habitats and Populations in the Northern Great Plains

Biotic response to the buildup of greenhouse gases in Earths atmosphere is considerably more complex than an adjustment to changing temperature and precipitation. The fertilization effect CO2 has on some plants, the impact UVB radiation has on health and productivity of organisms, and the resulting changes in competitive balance and trophic structure must also be considered. The intent of this paper is to review direct and indirect effects of anthropogenic greenhouse gases on wildlife, and to explore possible effects on populations of birds and their habitats in the northern Great Plains. Many of the potential effects of increasing greenhouse gases, such as declining plant nutritional value, changes in timing of insect emergence, and fewer and saltier wetlands, foreshadow a decline in avian populations on the Great Plains. However, other possible effects such as increased drought resistance and water use efficiency of vegetation, longer growing seasons, and greater overall plant biomass promise at least some mitigation. Effects of multiple simultaneous perturbations such as can be expected under doubled CO2 scenarios will require substantial basic research to clarify.