Targeted grazing reduces a widespread wetland plant invader with minimal nutrient impacts, yet native community recovery is limited.

Targeted grazing to control undesirable plant species is increasingly of interest across a diversity of ecosystems, particularly as an alternative or complement to widely used herbicides. However, there are limited comprehensive evaluations of targeted grazing that evaluate both invasive species management effectiveness and potential negative effects on the ecosystem. Phragmites australis, a tall-statured, dense perennial invasive grass from Eurasia, is a pervasive problem in wetlands across the North American continent. As with many invasive species where management has historically relied on herbicides and resistance is a growing concern, land managers seek viable alternatives that have minimal negative ecosystem impacts. Grazing has been used for millennia to manage native Phragmites in Europe. Similarly, in its invasive range within North America, small-scale studies suggest Phragmites may be suppressed by grazers. Yet, the effectiveness of grazing at large scales and its effects on broader ecosystem properties remain largely unknown. We evaluated the influence of targeted grazing on vegetation, soil nutrients, and water nutrients over two years in large plots (∼300x the size of previous studies). We also tested the effects of mowing, a treatment that can be used to facilitate grazer access to large, dense Phragmites stands. In line with our predictions, we found that cattle grazing effectively suppressed invasive Phragmites over two years. Mowing reduced litter, and moderately reduced standing dead Phragmites, both of which suppress native plant germination in this system. However, these reductions in Phragmites were not accompanied by indications of native plant community recovery, as we had optimistically predicted. Despite the potential for grazing to reduce nutrient sequestration by plants and fertilize soils, we were surprised to find no clear negative effects of grazing on nutrient mobilization to groundwater or floodwater. Taken together, our findings indicate that targeted grazing, when implemented at broad scales over short time frames, is effective at achieving invasive plant management goals without sizable nutrient impacts. However, additional steps will be needed to achieve the restoration of diverse, robust native plant communities.

Tipping the balance: The role of seed density, abiotic filters, and priority effects in seed-based wetland restoration.

Sowing native seeds is a common approach to reintroduce native plants to degraded systems. However, this method is often overlooked in wetland restoration despite the immense global loss of diverse native wetland vegetation. Developing guiding principles for seed-based wetland restoration is critical to maximize native plant recovery, particularly in previously invaded wetlands. Doing so requires a comprehensive understanding of how restoration manipulations, and their interactions, influence wetland plant community assembly. With a focus on the invader Phragmites australis, we established a series of mesocosm experiments to assess how native sowing density, invader propagule pressure, abiotic filters (water and nutrients), and native sowing timing (i.e., priority effects) interact to influence plant community cover and biomass in wetland habitats. Increasing the density of native seeds yielded higher native cover and biomass, but P. australis suppression with increasing sowing densities was minimal. Rather, community outcomes were largely driven by invader propagule pressure: P. australis densities of ≤500 seeds/m2 maintained high native cover and biomass. Low-water conditions increased the susceptibility of P. australis to dominance by native competitors. Early sowing of native seeds showed a large and significant benefit to native cover and biomass, regardless of native sowing density, suggesting that priority effects can be an effective restoration manipulation to enhance native plant establishment. Given the urgent wetland restoration need combined with the limited studies on seed-based wetland restoration, these findings provide guidance on restoration manipulations that are grounded in ecological theory to improve seed-based wetland restoration outcomes.

Metal concentrations in wetland plant tissues influences transfer to terrestrial food webs.

Wetland plants tolerate potentially hazardous metals through a variety of strategies, including exclusion or accumulation. Whether plants sequester metals and where they store them in their tissues is important for understanding the potential role of plants as remediators or vectors of metals to terrestrial food webs. Here we evaluate metal sequestration in Great Salt Lake wetlands for one invasive (Phragmites australis; phragmites) and three native plant species, i.e. threesquare bulrush (Schoenoplectus americanus), hardstem bulrush (Schoenoplectus acutus), alkali bulrush (Bolboschoenus maritimus), and their terrestrial invertebrates. We observed higher concentrations of arsenic and copper than other metals in plant tissues, although high lead concentrations were observed in phragmites. All plants acted as excluders of arsenic and selenium, retaining the bulk of the metal mass in belowground tissues. In contrast, lead, copper, and cadmium were transferred to above ground tissues of hardstem bulrush and phragmites. The aboveground translocation facilitated the movement of these metals into invertebrates, with the highest concentrations in most cases found in predators. Though our results highlight the potential for metal remediation via wetland plant growth and removal, care should be taken to ensure that remediation efforts do not lead to bioaccumulation.

The social-ecological system driving effective invasive plant management: two case studies of non-native Phragmites.

Globally, the management of invasive plants is motivated by a desire to improve ecosystem services (e.g., recreation, flood mitigation, soil fertility for agriculture, aesthetics) and critical habitat for imperiled species. To reduce invader populations and impacts, it is important to document the social and ecological basis (i.e., the social-ecological system) for the management that has been employed and areas where a greater level of coordination among stakeholder groups (managers, scientists, legislators, resource users) could improve efforts. We present a conceptual model that builds on current thinking for how best to connect these four stakeholder groups-to foster stronger citizen lobbying for impacted resources, science-based governance, legislator-driven noxious weed laws and funding for management and science, knowledge co-production by scientists and managers, and co-management by managers and resource users. In light of our model, we present two case studies based in Nebraska and Utah, U.S.A. involving a common North American wetland invader, Phragmites australis (non-native common reed). In Nebraska, potential lawsuits stemming from water conveyance was strong motivation for funding management. In Utah, duck hunters and other resource users initially instigated management. Progress toward the successful management of Phragmites has been the result of manager-scientist partnerships addressing a knowing-doing gap among practitioners, the complexities of management mosaics, as well as overcoming economic and logistical constraints. Our model demonstrates how legislative initiatives can fund new research and bolster on-going management, while organically building strong partnerships among scientists, managers, and resource users that are key for successfully managing invasive species.

Abiotic and Landscape Factors Constrain Restoration Outcomes Across Spatial Scales of a Widespread Invasive Plant.

The natural recolonization of native plant communities following invasive species management is notoriously challenging to predict, since outcomes can be contingent on a variety of factors including management decisions, abiotic factors, and landscape setting. The spatial scale at which the treatment is applied can also impact management outcomes, potentially influencing plant assembly processes and treatment success. Understanding the relative importance of each of these factors for plant community assembly can help managers prioritize patches where specific treatments are likely to be most successful. Here, using effects size analyses, we evaluate plant community responses following four invasive Phragmites australis management treatments (1: fall glyphosate herbicide spray, 2: summer glyphosate herbicide spray, 3: summer imazapyr herbicide spray, 4: untreated control) applied at two patch scales (12,000 m2 and 1,000 m2) and monitored for 5 years. Using variation partitioning, we then evaluated the independent and shared influence of patch scale, treatment type, abiotic factors, and landscape factors on plant community outcomes following herbicide treatments. We found that Phragmites reinvaded more quickly in large patches, particularly following summer herbicide treatments, while native plant cover and richness increased at a greater magnitude in small patches than large. Patch scale, in combination with abiotic and landscape factors, was the most important driver for most plant responses. Compared with the small plots, large patches commonly had deeper and more prolonged flooding, and were in areas with greater hydrologic disturbance in the landscape, factors associated with reduced native plant recruitment and greater Phragmites cover. Small patches were associated with less flooding and landscape disturbance, and more native plants in the surrounding landscape than large patches, factors which promoted higher native plant conservation values and greater native plant cover and richness. Herbicide type and timing accounted for very little of the variation in native plant recovery, emphasizing the greater importance of patch selection for better management outcomes. To maximize the success of treatment programs, practitioners should first manage Phragmites patches adjacent to native plant species and in areas with minimal hydrologic disturbance.

Invasive Phragmites australis management outcomes and native plant recovery are context dependent.

The outcomes of invasive plant removal efforts are influenced by management decisions, but are also contingent on the uncontrolled spatial and temporal context of management areas. Phragmites australis is an aggressive invader that is intensively managed in wetlands across North America. Treatment options have been understudied, and the ecological contingencies of management outcomes are poorly understood. We implemented a 5-year, multi-site experiment to evaluate six Phragmites management treatments that varied timing (summer or fall) and types of herbicide (glyphosate or imazapyr) along with mowing, plus a nonherbicide solarization treatment. We evaluated treatments for their influence on Phragmites and native plant cover and Phragmites inflorescence production. We assessed plant community trajectories and outcomes in the context of environmental factors. The summer mow, fall glyphosate spray treatment resulted in low Phragmites cover, high inflorescence reduction, and provided the best conditions for native plant recruitment. However, returning plant communities did not resemble reference sites, which were dominated by ecologically important perennial graminoids. Native plant recovery following initial Phragmites treatments was likely limited by the dense litter that resulted from mowing. After 5 years, Phragmites mortality and native plant recovery were highly variable across sites as driven by hydrology. Plots with higher soil moisture had greater reduction in Phragmites cover and more robust recruitment of natives compared with low moisture plots. This moisture effect may limit management options in semiarid regions vulnerable to water scarcity. We demonstrate the importance of replicating invasive species management experiments across sites so the contingencies of successes and failures can be better understood.

Strategic plant choices can alleviate climate change impacts: A review.

Ecosystem-based adaptation (EbA) uses biodiversity and ecosystem services to reduce climate change impacts to local communities. Because plants can alleviate the abiotic and biotic stresses of climate change, purposeful plant choices could improve adaptation. However, there has been no systematic review of how plants can be applied to alleviate effects of climate change. Here we describe how plants can modify climate change effects by altering biological and physical processes. Plant effects range from increasing soil stabilization to reducing the impact of flooding and storm surges. Given the global scale of plant-related activities such as farming, landscaping, forestry, conservation, and restoration, plants can be selected strategically-i.e., planting and maintaining particular species with desired impacts-to simultaneously restore degraded ecosystems, conserve ecosystem function, and help alleviate effects of climate change. Plants are a tool for EbA that should be more broadly and strategically utilized.

Cosmopolitan Species As Models for Ecophysiological Responses to Global Change: The Common Reed Phragmites australis.

Phragmites australis is a cosmopolitan grass and often the dominant species in the ecosystems it inhabits. Due to high intraspecific diversity and phenotypic plasticity, P. australis has an extensive ecological amplitude and a great capacity to acclimate to adverse environmental conditions; it can therefore offer valuable insights into plant responses to global change. Here we review the ecology and ecophysiology of prominent P. australis lineages and their responses to multiple forms of global change. Key findings of our review are that: (1) P. australis lineages are well-adapted to regions of their phylogeographic origin and therefore respond differently to changes in climatic conditions such as temperature or atmospheric CO2; (2) each lineage consists of populations that may occur in geographically different habitats and contain multiple genotypes; (3) the phenotypic plasticity of functional and fitness-related traits of a genotype determine the responses to global change factors; (4) genotypes with high plasticity to environmental drivers may acclimate or even vastly expand their ranges, genotypes of medium plasticity must acclimate or experience range-shifts, and those with low plasticity may face local extinction; (5) responses to ancillary types of global change, like shifting levels of soil salinity, flooding, and drought, are not consistent within lineages and depend on adaptation of individual genotypes. These patterns suggest that the diverse lineages of P. australis will undergo intense selective pressure in the face of global change such that the distributions and interactions of co-occurring lineages, as well as those of genotypes within-lineages, are very likely to be altered. We propose that the strong latitudinal clines within and between P. australis lineages can be a useful tool for predicting plant responses to climate change in general and present a conceptual framework for using P. australis lineages to predict plant responses to global change and its consequences.

Biotic resistance, disturbance, and mode of colonization impact the invasion of a widespread, introduced wetland grass.

Disturbance and biotic resistance are important factors driving plant invasions, but how these factors interact for plants with different modes of colonization (i.e., sexual and asexual) is unclear. We evaluated factors influencing the invasion of nonnative Phragmites australis, which has been rapidly expanding in brackish tidal wetlands in Chesapeake Bay. We conducted a survey of naturally occurring small-scale disturbances (removal of vegetation and/or sediment deposition) across four plant communities; determined the effects of small-scale disturbance and biotic resistance on P. australis seedling and rhizome emergence; and tested the effects of size and frequency of small-scale disturbances on seedling emergence and survival of transplanted seedlings. The results of our study demonstrate that the invasion window for seeds is in disturbed areas in high-marsh plant communities that flood less frequently; seedling emergence in undisturbed areas was negligible. Establishment of shoots from rhizome segments was low in all plant communities. Disturbance size and frequency had no significant impact on seed germination and seedling survival. Our findings provide evidence that small-scale within-wetland disturbances are important for the invasion of the nonnative lineage of P. australis by seeds in brackish tidal wetlands in Chesapeake Bay. Efforts to reduce disturbances, large and small, in wetlands can be used to limit P. australis invasion by seed, but invasion by rhizome is still likely to occur across many plant communities irrespective of the presence of disturbance.

Phragmites australis management in the United States: 40 years of methods and outcomes.

Studies on invasive plant management are often short in duration and limited in the methods tested, and lack an adequate description of plant communities that replace the invader following removal. Here we present a comprehensive review of management studies on a single species, in an effort to elucidate future directions for research in invasive plant management. We reviewed the literature on Phragmites management in North America in an effort to synthesize our understanding of management efforts, identify gaps in knowledge and improve the efficacy of management. Additionally, we assessed recent ecological findings concerning Phragmites mechanisms of invasion and integrated these findings into our recommendations for more effective management. Our overall goal is to examine whether or not current management approaches can be improved and whether they promote reestablishment of native plant communities. We found: (i) little information on community-level recovery of vegetation following removal of Phragmites; and (ii) most management approaches focus on the removal of Phragmites from individual stands or groups of stands over a relatively small area. With a few exceptions, recovery studies did not monitor vegetation for substantial durations, thus limiting adequate evaluation of the recovery trajectory. We also found that none of the recovery studies were conducted in a landscape context, even though it is now well documented that land-use patterns on adjacent habitats influence the structure and function of wetlands, including the expansion of Phragmites. We suggest that Phragmites management needs to shift to watershed-scale efforts in coastal regions, or larger management units inland. In addition, management efforts should focus on restoring native plant communities, rather than simply eradicating Phragmites stands. Wetlands and watersheds should be prioritized to identify ecosystems that would benefit most from Phragmites management and those where the negative impact of management would be minimal.

Moving from a regional to a continental perspective of Phragmites australis invasion in North America.

AIMS: We use a regional comparison of Phragmites australis (common reed) subsp. americanus, P. australis subsp. berlandieri and introduced P. australis (possibly five sublineages) in the Chesapeake Bay, the St Lawrence River, Utah and the Gulf Coast to inform a North American perspective on P. australis invasion patterns, drivers, impacts and research needs. FINDINGS AND RESEARCH NEEDS: Our regional assessments reveal substantial diversity within and between the three main lineages of P. australis in terms of mode of reproduction and the types of environment occupied. For introduced P. australis, the timing of introduction also differed between the regions. Nevertheless, a common finding in these regions reinforces the notion that introduced P. australis is opportunistic and thrives in disturbed habitats. Thus, we expect to see substantial expansion of introduced P. australis with increasing anthropogenic disturbances in each of these regions. Although there have been some studies documenting the negative impacts of introduced P. australis, it also plays a beneficial role in some regions, and in some cases, the purported negative impacts are unproven. There is also a broader need to clarify the genetic and ecological relationships between the different introduced sublineages observed in North America, and their relative competitive ability and potential for admixture. This may be done through regional studies that use similar methodologies and share results to uncover common patterns and processes. To our knowledge, such studies have not been performed on P. australis in spite of the broad attention given to this species. Such research could advance theoretical knowledge on biological invasion by helping to determine the extent to which the patterns observed can be generalized or are sublineage specific or region specific. SYNTHESIS: Given what appears to be sometimes idiosyncratic invasion patterns when interpreted in isolation in the regions that we analysed, it may be time to consider initiatives on a continental (if not intercontinental) scale to tackle unresolved issues about P. australis.

Isolation of 11 polymorphic tri- and tetranucleotide microsatellite loci in a North American sedge (Carex scoparia: Cyperaceae) and cross-species amplification in three additional Carex species.

We report on the isolation and evaluation of 11 microsatellites from a widespread eastern North American wetland sedge, Carex scoparia. Loci exhibit 3-9 alleles over five populations and significant F(IS) (0.204-0.717) in most populations. All primers cross-amplify in at least two other species, and 10 cross-amplify in the more distantly related C. stipata. These markers will be used to examine population genetics and patterns of chromosomal diversification in this ecologically important sedge species and its relatives.

Effect of light on seed germination of eight wetland Carex species.

BACKGROUND AND AIMS: In wetland plant communities, species-specific responses to pulses of white light and to red : far-red light ratios can vary widely and influence plant emergence from the seed bank. Carex species are the characteristic plants of sedge meadows of natural prairie wetlands in mid-continental USA but are not returning to restored wetlands. Little is known about how light affects seed germination in these species-information which is necessary to predict seed bank emergence and to develop optimal revegetation practices. The effects of light on germination in eight Carex species from prairie wetlands were investigated. METHODS: Non-dormant seeds of eight Carex species were used to determine the influence of light on germination by examining: (a) the ability of Carex seeds to germinate in the dark; (b) the effect of different lengths of exposures to white light on germination; (c) whether the effect of white light can be replaced by red light; and (d) whether the germination response of Carex seeds to white or red light is photoreversible by far-red light. KEY RESULTS: Seeds of C. brevior and C. stipata germinated >25 % in continuous darkness. Germination responses after exposure to different lengths of white light varied widely across the eight species. Carex brevior required <15 min of white light for > or =50 % germination, while C. hystericina, C. comosa, C. granularis and C. vulpinoidea required > or =8 h. The effect of white light was replaced by red light in all species. The induction of germination after exposure to white or red light was reversed by far-red light in all species, except C. stipata. CONCLUSIONS: The species-specific responses to simulated field light conditions suggest that (a) the light requirements for germination contribute to the formation of persistent seed banks in these species and (b) in revegetation efforts, timing seed sowing to plant community development and avoiding cover crops will improve Carex seed germination.