Exotic Annuals Research Articles

Nutrient versus seed bank depletion approaches to controlling exotic annuals in threatened Box Gum woodlands

AbstractControl of exotic annuals is often a priority when restoring degraded grasslands or shrublands. This study evaluated combinations of nutrient‐depleting (carbon addition) and seed bank‐depleting approaches for controlling exotic annuals, and compared the seed bank depletion technique of spring burning with the more easily applied technique of pulse grazing. Treatments were applied in two Box Gum woodlands over 4 years. Consistent with earlier studies, carbon addition dramatically reduced exotic annuals and available nutrients in all 3 years at both sites. Exotic annual grass abundance was significantly reduced in burnt plots following the first year and in grazed plots following the second year of application. Spring burning or grazing did not reduce available nutrients or exotic annual broadleaf abundance at either site. The effect of carbon addition on exotic annuals and available nutrients was so powerful that no additional benefit of the combination treatments was found, although at one site burning and grazing slightly reduced the effectiveness of carbon addition in suppressing broadleaf exotic annuals. We conclude that nutrient‐depleting approaches are most effective because they control both exotic annual grasses and broadleaf exotics annuals, but given their expense seed bank‐depleting approaches may be more practical where exotic annual grasses dominate. In particular, pulse grazing is readily applied across large areas, offering a relatively simple tool that if appropriately implemented could enhance outcomes of restoration investments in vegetation communities invaded by exotic annual grasses.

Short-term priority over exotic annuals increases the initial density and longer-term cover of native perennial grasses.

Temporal priority can affect individual performance and reproduction, as well as community assembly, but whether these effects persist over time remains unclear, and their demographic mechanisms have been little explored. The continued dominance of exotic annual grasses in California has been commonly attributed to their demonstrated early germination and rapid early growth relative to native perennial grasses. This advantage may play a crucial role in the structure of California exotic annual grasslands, as well as in the practice of native grassland restoration. We tested whether a two-week planting advantage under field conditions increased individual survival, growth, and reproduction for four native perennial grass species and whether these effects persisted over three years. We show that short-term priority significantly increased the establishment success of' native perennial grasses. Increased density of native grass seedlings presaged later large increases in cover that were not evident in the first year after planting. Although priority effects at the individual level may diminish over time, short differences in emergence timing can have long-lasting effects on community structure. Earlier germination and faster initial growth of exotic annual species may help explain their unprecedented invasion and continued dominance of California grasslands. Finally, these results highlight the importance of priority effects for effective exotic annual control during native grassland restoration in California: initial control can increase the establishment of native perennial seedlings, which then results in long-term control by mature native individuals.

Extractable nitrogen and microbial community structure respond to grassland restoration regardless of historical context and soil composition.

Grasslands have a long history of invasion by exotic annuals, which may alter microbial communities and nutrient cycling through changes in litter quality and biomass turnover rates. We compared plant community composition, soil chemical and microbial community composition, potential soil respiration and nitrogen (N) turnover rates between invaded and restored plots in inland and coastal grasslands. Restoration increased microbial biomass and fungal : bacterial (F : B) ratios, but sampling season had a greater influence on the F : B ratio than did restoration. Microbial community composition assessed by phospholipid fatty acid was altered by restoration, but also varied by season and by site. Total soil carbon (C) and N and potential soil respiration did not differ between treatments, but N mineralization decreased while extractable nitrate and nitrification and N immobilization rate increased in restored compared with unrestored sites. The differences in soil chemistry and microbial community composition between unrestored and restored sites indicate that these soils are responsive, and therefore not resistant to feedbacks caused by changes in vegetation type. The resilience, or recovery, of these soils is difficult to assess in the absence of uninvaded control grasslands. However, the rapid changes in microbial and N cycling characteristics following removal of invasives in both grassland sites suggest that the soils are resilient to invasion. The lack of change in total C and N pools may provide a buffer that promotes resilience of labile pools and microbial community structure.

Initial success of native grasses is contingent on multiple interactions among exotic grass competition, temporal priority, rainfall and site effects.

Ecological communities are increasingly being recognized as the products of contemporary drivers and historical legacies that are both biotic and abiotic. In an attempt to unravel multiple layers of ecological contingency, we manipulated (i) competition with exotic annual grasses, (ii) the timing of this competition (temporal priority in arrival/seeding times) and (iii) watering (simulated rainfall) in a restoration-style planting of native perennial grasses. In addition, we replicated this experiment simultaneously at three sites in north-central California. Native perennial grasses had 73-99 % less cover when planted with exotic annuals than when planted alone, but this reduction was greatly ameliorated by planting the natives 2 weeks prior to the exotics. In a drought year, irrigation significantly reduced benefits of early planting so that these benefits resembled those observed in a non-drought year. There were significant differences across the three sites (site effects and interactions) in (i) overall native cover, (ii) the response of natives to competition, (iii) the strength of the temporal priority effect and (iv) the degree to which supplemental watering reduced priority effects. These results reveal the strong multi-layered contingency that underlies even relatively simple communities.

Remote sensing analysis of vegetation recovery following short-interval fires in Southern California shrublands.

Increased fire frequency has been shown to promote alien plant invasions in the western United States, resulting in persistent vegetation type change. Short interval fires are widely considered to be detrimental to reestablishment of shrub species in southern California chaparral, facilitating the invasion of exotic annuals and producing “type conversion”. However, supporting evidence for type conversion has largely been at local, site scales and over short post-fire time scales. Type conversion has not been shown to be persistent or widespread in chaparral, and past range improvement studies present evidence that chaparral type conversion may be difficult and a relatively rare phenomenon across the landscape. With the aid of remote sensing data covering coastal southern California and a historical wildfire dataset, the effects of short interval fires (<8 years) on chaparral recovery were evaluated by comparing areas that burned twice to adjacent areas burned only once. Twelve pairs of once- and twice-burned areas were compared using normalized burn ratio (NBR) distributions. Correlations between measures of recovery and explanatory factors (fire history, climate and elevation) were analyzed by linear regression. Reduced vegetation cover was found in some lower elevation areas that were burned twice in short interval fires, where non-sprouting species are more common. However, extensive type conversion of chaparral to grassland was not evident in this study. Most variables, with the exception of elevation, were moderately or poorly correlated with differences in vegetation recovery.

Soil Resources Influence Vegetation and Response to Fire and Fire-Surrogate Treatments in Sagebrush-Steppe Ecosystems

Current paradigm suggests that spatial and temporal competition for resources limit an exotic invader, cheatgrass (Bromus tectorum L.), which once established, alters fire regimes and can result in annual grass dominance in sagebrush steppe. Prescribed fire and fire surrogate treatments (mowing, tebuthiuron, and imazapic) are used to reduce woody fuels and increase resistance to exotic annuals, but may alter resource availability and inadvertently favor invasive species. We used four study sites within the Sagebrush Steppe Treatment Evaluation Project (SageSTEP) to evaluate 1) how vegetation and soil resources were affected by treatment, and 2) how soil resources influenced native herbaceous perennial and exotic annual grass cover before and following treatment. Treatments increased resin exchangeable NH4+, NO3-, H2PO4-, and K+, with the largest increases caused by prescribed fire and prolonged by application of imazapic. Burning with imazapic application also increased the number of wet growing degree days. Tebuthiuron and imazapic reduced exotic annual grass cover, but imazapic also reduced herbaceous perennial cover when used with prescribed fire. Native perennial herbaceous species cover was higher where mean annual precipitation and soil water resources were relatively high. Exotic annual grass cover was higher where resin exchangeable H2PO4- was high and gaps between perennial plants were large. Prescribed fire, mowing, and tebuthiuron were successful at increasing perennial herbaceous cover, but the results were often ephemeral and inconsistent among sites. Locations with sandy soil, low mean annual precipitation, or low soil water holding capacity were more likely to experience increased exotic annual grass cover after treatment, and treatments that result in slow release of resources are needed on these sites. This is one of few studies that correlate abiotic variables to native and exotic species cover across a broad geographic setting, and that demonstrates how soil resources potentially influence the outcome of management treatments.

Plant Recruitment and Soil Microbial Characteristics of Rehabilitation Seedings Following Wildfire in Northern Utah

AbstractOne goal of post‐fire native species seeding is to increase plant community resistance to exotic weed invasions, yet few studies address the impacts of seeding on exotic annual establishment and persistence. In 2010 and 2011, we investigated the influence of seedings on exotic annuals and the underlying microbial communities. The wildfire site in northern Utah was formerly dominated by Artemisia tridentata ssp. wyomingensis, but burned in September 2008. Experimental seeding treatments were installed in November 2008 to examine strategies for establishing native species using two drills, hand broadcasts and different timing of seed applications (resulting in 13 seeding treatments). We collected aboveground biomass of invasive annuals (Halogeton glomeratus, Salsola kali, and Bromus tectorum), other volunteer plants from the extant seed bank, and seeded species from all treatments in the second and third years after fire. We sampled soils within microsites beneath native perennial bunchgrass and exotic annuals to characterize underlying soil microbial communities. High precipitation following seeding led to strong seedling establishment and we found few differences between seeding treatments established with either drill. All seeded treatments reduced exotic biomass by at least 90% relative to unseeded controls. Soil microbial communities (phospholipid fatty acid analysis), beneath B. tectorum, Poa secunda, and Pseudoroegneria spicata microsites differed little 3 years after fire. However, microbial abundance beneath P. spicata increased from June to July, suggesting that microbial communities beneath successful seedings can vary greatly within a single growing season.

Attempting to Restore Herbaceous Understories in Wyoming Big Sagebrush Communities with Mowing and Seeding

AbstractShrub steppe communities with depleted perennial herbaceous understories often need to be restored to increase resilience and resistance. Mowing has been applied to Wyoming big sagebrush (Artemisia tridentata Nutt. ssp. wyomingensis Beetle &amp; Young) steppe plant communities to reduce sagebrush dominance and restore native herbaceous vegetation, but success has been limited and hampered by increases in exotic annuals. Seeding native bunchgrasses after mowing may accelerate recovery and limit exotics. We compared mowing followed by drill‐seeding native bunchgrasses to mowing and an untreated control at five sites in southeastern Oregon over a 4‐year period. Mowing and seeding bunchgrasses increased bunchgrass density; however, bunchgrass cover did not differ among treatments. Exotic annuals increased with mowing whether or not post‐mowing seeding occurred. Mowing, whether or not seeding occurred, also reduced biological soil crusts. Longer term evaluation is needed to determine if seeded bunchgrasses will increase enough to suppress exotic annuals. Seeded bunchgrasses may have been limited by increases in exotic annuals. Though restoration of sagebrush communities with degraded understories is needed, we do not recommend mowing and seeding native bunchgrasses because this treatment produced mixed results that may lower the resilience and resistance of these communities. Before this method is applied, research is needed to increase our understanding of how to improve establishment of seeded native bunchgrasses. Alternatively, restoration practitioners may need to apply treatments to control exotic annuals and repeatedly seed native bunchgrasses.

Restoring the Sagebrush Component in Crested Wheatgrass–Dominated Communities

Monotypic stands of crested wheatgrass (Agropyron cristatum [L] Gaertm. and Agropyron desertorum [Fisch.] Schult.), an introduced grass, occupy vast expanses of the sagebrush steppe. Efforts to improve habitat for sagebrush-associated wildlife by establishing a diverse community of native vegetation in crested wheatgrass stands have largely failed. Instead of concentrating on a diversity of species, we evaluated the potential to restore the foundation species, Wyoming big sagebrush (Artemisia tridentata spp. wyomingensis [Beetle & A. Young] S. L. Welsh), to these communities. We investigated the establishment of Wyoming big sagebrush into six crested wheatgrass stands (sites) by broadcast seeding and planting seedling sagebrush across varying levels of crested wheatgrass control with glyphosate. Planted sagebrush seedlings survived at high rates (∼ 70% planted sagebrush survival 3yr postplanting), even without crested wheatgrass control. However, most attempts to establish sagebrush by broadcast seeding failed. Only at high levels of crested wheatgrass control did a few sagebrush plants establish from broadcasted seed. Sagebrush density and cover were greater with planting seedlings than broadcast seeding. Sagebrush cover, height, and canopy area were greater at higher levels of crested wheatgrass control. High levels of crested wheatgrass control also created an opportunity for exotic annuals to increase. Crested wheatgrass rapidly recovered after glyphosate control treatments, which suggests multiple treatments may be needed to effectively control crested wheatgrass. Our results suggest that planting sagebrush seedlings can structurally diversify monotypic crested wheatgrass stands to provide habitat for sagebrush-associated wildlife. Though this is not the full diversity of native functional groups representative of the sagebrush steppe, it is a substantial improvement over other efforts that have largely failed to alter these plant communities. We also hypothesize that planting sagebrush seedlings in patches or strips may provide a relatively inexpensive method to facilitate sagebrush recovery across vast landscapes where sagebrush has been lost.

Annual-perennial plant relationships and species selection for desert restoration

Exotic plant invasion is a growing concern in the conservation and management of indigenous arid land ecosystems. By creating areas of ameliorated microclimates and fertile soil below their canopies, perennial plants might influence exotic annual plant invasions. We conducted a quantitative literature review of studies that compared exotic annual plant abundance among native perennial plant species and interspace (open areas) microsites in North America’s Mojave Desert, where exotic plant invasion has corresponded with increasing extent of wildfire and broad-scale ecosystem transformation. Ten studies compared exotic annual plant abundance between interspaces and below a total of 36 native perennial species. These studies revealed that: (1) With few exceptions, most native perennial species supported a greater abundance of exotic annuals than interspaces, indicating overall facilitation of exotic species by native perennials. (2) Exotic species abundance varied by orders of magnitude among native perennial species, with some perennial species harboring amounts of exotics similar to interspaces. (3) Distributions of dominant exotic species varied, where Bromus rubens displayed a greater affinity for below-perennial microsites than did Schismus spp. and Erodium cicutarium that often were most abundant in interspaces. Results suggest that the degree of facilitation of exotic plants warrants consideration when selecting native perennial species for revegetation and restoration projects.

Relationship between habitat characteristics and densities of southern Idaho ground squirrels

Many wildlife species living in sagebrush-steppe habitats of the Intermountain West of the United States have experienced population declines. Effective management of such species, especially restoration efforts, is contingent on understanding relationships between habitat characteristics and population densities of the target species. Unfortunately, even such basic information is often lacking. The goal of this 2-year study was to determine the relationship between southern Idaho ground squirrel (Urocitellus endemicus) population densities and habitat variables: soils (texture), topography (slope, aspect), and vegetation (canopy cover, species diversity). We measured population density indirectly through burrow entrance counts and categorized them into high-burrow and low-burrow densities. We used logistic regression and Akaike's Information Criterion to identify a best subset of models. We employed model averaging and calculated odds ratios for averaged parameter estimates found in the best models. A high density of burrows was associated with greater percentages of silt; east-facing aspects; greater plant species diversity; and greater cover of perennial grasses, perennial grasses and forbs, and native perennial forbs. Low burrow density was associated with greater percentages of sand; south-facing aspects; greater cover of exotic annuals; and lesser plant species diversity. Management of southern Idaho ground squirrel habitat should focus on protecting areas with existing native vegetation and restoring native, perennial vegetation in areas that are infested with exotic annuals, especially in areas possessing suitable soil types and topographic features. © 2013 The Wildlife Society.

Estimating Winter Annual Biomass in the Sonoran and Mojave Deserts with Satellite- and Ground-Based Observations

Winter annual plants in southwestern North America influence fire regimes, provide forage, and help prevent erosion. Exotic annuals may also threaten native species. Monitoring winter annuals is difficult because of their ephemeral nature, making the development of a satellite monitoring tool valuable. We mapped winter annual aboveground biomass in the Desert Southwest from satellite observations, evaluating 18 algorithms using time-series vegetation indices (VI). Field-based biomass estimates were used to calibrate and evaluate each algorithm. Winter annual biomass was best estimated by calculating a base VI across the period of record and subtracting it from the peak VI for each winter season (R2 = 0.92). The normalized difference vegetation index (NDVI) derived from 8-day reflectance data provided the best estimate of winter annual biomass. It is important to account for the timing of peak vegetation when relating field-based estimates to satellite VI data, since post-peak field estimates may indicate senescent biomass which is inaccurately represented by VI-based estimates. Images generated from the best-performing algorithm show both spatial and temporal variation in winter annual biomass. Efforts to manage this variable resource would be enhanced by a tool that allows the monitoring of changes in winter annual resources over time.

A Landscape Similarity Index: Multitemporal Remote Sensing to Track Changes in Big Sagebrush Ecological Sites

A similarity index for big sagebrush ecological sites was developed in northern Utah. In contrast to field measurements used to calculate similarity to reference states, our approach relies on the utilization of historic archives of satellite imagery to measure the ecological distance to benchmarks of undesired conditions such as invasion by exotic annuals and woodland encroachment. Our benchmarks consisted of locations for which there are field data collected for monitoring and evaluation purposes for several time periods. We utilized a temporal series of Landsat thematic mapper (TM) imagery that spanned 1984 to 2008 from which the soil-adjusted vegetation index (SAVI) and other transformations were extracted. Topographic and climatic variables were also included as ancillary data. Multidimensional scaling (MDS) was used to obtain scores in reduced ordination space for two periods of interest: 1984–1996 and 1997–2008. Interannual SAVI mean-variance plots provided evidence that the benchmarks and ecological sites have a distinct temporal response that allows an objective comparison. Our MDS results also show that natural clusters can be identified in the reduced statistical space for ecological sites that are a dominant component of a soil map unit. The two MDS solutions allowed the ordination of ecological sites in two gradients of productivity and bare ground. Interpretations of the transitions and trajectories of mountain, Wyoming, and basin big sagebrush sites correlated well with the ecological expectation. We anticipate that range conservationists and others actively working in rangeland evaluation can use this application to develop and update ecological site descriptions and state-and-transition models from a remotely sensed perspective.

Manipulating livestock grazing to enhance native plant diversity and cover in native grasslands

Temperate perennial grasslands globally have been subject to extensive biodiversity loss. Identifying livestock grazing regimes that maintain and enhance the diversity and cover of native plant species in these ecosystems remains a key challenge. The responses of vegetation to different sheep grazing regimes were assessed over 3 years in grasslands of south-eastern Australia. An open communal experimental design was used to assess the effects of varying season and duration of exclusion of grazing by sheep, replicated at three locations. Manipulation of season and duration of exclusion of grazing led to few major changes in the cover of native perennial grasses or forbs, although seasonal variation was considerable. Exclusion of grazing in the spring did increase the likelihood of occurrence of grazing-sensitive native forb species but also lead to an increase in the cover of exotic annual species. However, cover of exotic annual species tended to decline with increasing duration of exclusion, while the abundance of native, grazing-sensitive forbs and the cover of perennial grasses increased. Small-scale richness of native perennial forb species increased with a 3-month period of exclusion of grazing, but declined with year-round exclusion of sheep. Total species richness also declined in response to year-round exclusion of sheep and rates of decline were correlated with the rate at which herbage mass accumulated. While strategic grazing did not result in major vegetation changes in the short term, it is suggested that some grazing exclusion may enhance the survival of infrequent species most sensitive to sheep grazing. Caution, however, should be taken when grazing regimes implemented benefit both desirable (native forbs) and undesirable (exotic annuals) species. Having a mosaic of flexible grazing management regimes across the landscape is likely to be beneficial for native plant diversity.

Exotic annuals reduce soil heterogeneity in coastal sage scrub soil chemical and biological characteristics

Exotic plant invasions alter ecosystem structure and function above- and below-ground through plant–soil feedbacks. The resistance of ecosystems to invasion can be measured by the degree of change in microbial communities and soil chemical pools and fluxes, whereas their resilience can be measured by the ability to recover following restoration. Coastal sage scrub (CSS) is one of the most highly invaded ecosystems in the US but the response of CSS soils to exotic plant invasion is little known. We examined resistance and resilience of CSS soil chemical and biological characteristics following invasion of exotic annual grasses and forbs and restoration of the native plant community. We hypothesized that invasion of exotic plant species would change biological and chemical characteristics of CSS soils by altering soil nutrient inputs. Additionally, we expected that if exotic plants were controlled and native plants were restored, native soil characteristics would recover. We sampled two locations with invaded, restored and native CSS for plant community composition, soil chemistry and microbial communities, and phospholipid fatty acid (PLFA) profiles. Communities invaded by exotic annuals were resistant to some measured parameters but not others. Extractable nitrogen pools decreased, nitrogen cycling rates increased, and microbial biomass and fungal:bacterial ratios were altered in invaded soils, and these effects were mediated by the phenological stage of the dominant plant species. The largest impact of invasion on soils was an overall reduction of spatial heterogeneity in soil nutrients, nutrient cycling and microbial communities. Restored plots tended to recover in most biotic and chemical parameters including increased resource heterogeneity compared to invaded plots, suggesting that CSS soils are resilient but not resistant to invasion.

Mowing Wyoming Big Sagebrush Communities With Degraded Herbaceous Understories: Has a Threshold Been Crossed?

Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis &lsqb;Beetle & A. Young] S.L. Welsh) plant communities with degraded native herbaceous understories occupy vast expanses of the western United States. Restoring the native herbaceous understory in these communities is needed to provide higher-quality wildlife habitat, decrease the risk of exotic plant invasion, and increase forage for livestock. Though mowing is commonly applied in sagebrush communities with the objective of increasing native herbaceous vegetation, vegetation response to this treatment in degraded Wyoming big sagebrush communities is largely unknown. We compared mowed and untreated control plots in five Wyoming big sagebrush plant communities with degraded herbaceous understories in eastern Oregon for 3 yr posttreatment. Native perennial herbaceous vegetation did not respond to mowing, but exotic annuals increased with mowing. Density of cheatgrass (Bromus tectorum L.), a problematic exotic annual grass, was 3.3-fold greater in the mowed than untreated control treatment in the third year posttreatment. Annual forb cover, largely consisting of exotic species, was 1.8-fold greater in the mowed treatment compared to the untreated control in the third year posttreatment. Large perennial grass cover was not influenced by mowing and remained below 2%. Mowing does not appear to promote native herbaceous vegetation in degraded Wyoming big sagebrush plant communities and may facilitate the conversion of shrublands to exotic annual grasslands. The results of this study suggest that mowing, as a stand-alone treatment, does not restore the herbaceous understory in degraded Wyoming big sagebrush plant communities. We recommend that mowing not be applied in Wyoming big sagebrush plant communities with degraded understories without additional treatments to limit exotic annuals and promote perennial herbaceous vegetation. Las comunidades de plantas de artemisia Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis &lsqb;Beetle & A. Young] S.L. Welsh) con degradadas coberturas herbáceas ocupan una gran extensión del oeste de los Estados Unidos. El restablecimiento de la cobertura herbácea nativa en estas comunidades es necesario para mejorar la calidad del hábitat para fauna silvestre, mitigar el riesgo de la invasión de plantas exóticas e incrementar la producción de forraje para ganado. A pesar de que comúnmente se hacen cortes en comunidades de artemisia con el objetivo de incrementar la vegetación nativa, se desconoce la respuesta de la vegetación al tratamiento en áreas con comunidades degradadas de Wyoming big sagebrush. Se compararon parcelas segadas y áreas control sin tratamientos en cinco comunidades de Wyoming big sagebrush con cobertura herbácea degradada en el este de Oregón durante tres años posteriores a la aplicación de los tratamientos. La vegetación perenne herbácea no respondió a la siega, pero las plantas exóticas anuales se incrementaron con esta práctica. La densidad de cheatgrass (Bromus tectorum L.), una especie problemática exótica fue tres veces mayor en áreas segadas que en áreas control sin tratamiento tres años después de la aplicación de los tratamientos. La cobertura herbácea anual en gran parte formada por especies exóticas fue 1.8 veces mayor en las zonas segadas que en las áreas control sin tratamiento tres años posteriores a la aplicación de tratamientos. La gran cobertura de pastos perennes no fue influenciada por la siega y permaneció debajo del 2%. La siega parece no promover la vegetación herbácea nativa en comunidades degradadas de Wyoming big sagebrush y podría facilitar el cambio de áreas de matorrales a pastizales anuales exóticos. Los resultados de este estudio sugieren que la siega, como un tratamiento independiente, no restablece la cobertura herbácea en comunidades de Wyoming big sagebrush. Nosotros recomendamos que la siega no se practique en comunidades de Wyoming big sagebrush con cobertura degradada sin tratamientos adicionales para limitar la presencia de plantas anuales exóticas y estimular la vegetación herbácea perenne.

Effects of flood timing and livestock grazing on exotic annual plants in riverine floodplains

Summary Riverine floodplains around the world are highly prone to invasions by exotic species. While many processes have contributed to floodplain invasions, two theories have received particular attention: (i) replacement of native perennials by exotic annuals following historical livestock grazing and (ii) ‘terrestrialization’ because of fewer floods following river regulation. We assessed how exclusion of grazers and flooding in different seasons affected established populations of exotic annuals in an 8‐year experiment. We hypothesized that effects of season of flooding would vary depending on timing in relation to annual plant life cycles and that outcomes may be affected by interactions between grazing and flooding. Flooding in any season reduced exotic annuals to &lt;2% cover and converted a dryland flora dominated by exotic annuals to a wetland flora dominated by native perennials. However, flood effects may be short lived, as areas flooded in 1 year supported abundant exotics the next. Grazer exclusion and interactions between grazing and flooding did not affect cover of exotic annuals, although grazing depleted native perennials in recently flooded areas. Synthesis and applications. Consistent with the terrestrialization hypothesis, short‐duration floods provide a practical method for controlling exotic annual plants where river regulation has reduced the frequency of natural floods. As exotic impacts are greatest in dry periods, when exotics are most abundant, ecosystem benefits will be greatest when flooding leads to a protracted decline in exotic annuals during the dry (non‐inundated) years following flood recession. Otherwise, reductions in exotic annuals may be transient and have little positive impact on floodplain ecosystems and processes.

MORE EXOTIC AND FEWER NATIVE PLANT SPECIES: RIVERINE VEGETATION PATTERNS ASSOCIATED WITH ALTERED SEASONAL FLOW PATTERNS

ABSTRACTNatural flow regimes are important for sustaining riverine vegetation. The regulation of river flows to provide water for agriculture often results in changes to flow timing. This study assesses the impact of altered seasonal flow patterns on riverine flora.Within temperate Australia, we surveyed the vegetation of five lowland rivers, three of which have large dams that alter their seasonal flow patterns; the other two are unregulated. From four to six sites were selected on each river, and these were classified into three levels of regulation based on the extent to which the timing of their seasonal flow patterns were altered. Sites were surveyed in winter and the following summer. Permanent quadrats were also established at a number of the surveyed sites and resurveyed every 3 months.Of the 267 plant taxa identified, 145 were exotic (non‐native). More exotic taxa and fewer native taxa were associated with increasing level of seasonal flow inversion (regulation). In particular, greater numbers of short‐lived exotic terrestrial taxa and fewer native woody taxa were associated with increasing level of regulation. Some exotic woody species (e.g. willows) were more common in the unregulated rivers and may have life‐history traits favoured by the natural seasonal flow patterns of study area. Multivariate analyses showed that level of regulation had a significant effect on the overall composition of the riverine vegetation.Our results provide support for the hypotheses that flow regulation adversely affects native species diversity and increases the vulnerability of riparian zones to invasion by exotic species; however, these effects are dependent on plant species' life‐history strategies. Our study highlights the importance of natural seasonal flow patterns for sustaining native riverine plant communities. Flow management aimed at maintaining or restoring ecological values should consider seasonal flow patterns. Winter/spring flow peaks may be particularly important for the recruitment of native riverine plants, especially trees and shrubs, and reducing the extent of exotic annuals and grasses. Copyright © 2012 John Wiley &amp; Sons, Ltd.

Relationships among soil fertility, native plant diversity and exotic plant abundance inform restoration of forb‐rich eucalypt woodlands

AbstractAim Water and nutrient availability are major limits to productivity in semi‐arid ecosystems; hence, ecological restoration often focuses on conserving or concentrating soil resources. By contrast, nutrient enrichment can promote invasion by exotic annuals, leading to restoration approaches that target reduction of soil nutrients. We aimed to explore potential biodiversity trade‐offs between these approaches by investigating relationships among soil nutrients, exotic annuals and native plant diversity and composition. In particular, we investigated the hypothesis that native plant diversity in semi‐arid to temperate woodlands reflects the productivity–diversity hypothesis, leading to hump‐backed relationships with soil nutrients such that (1) native plant diversity declines with increasing nutrient enrichment and (2) native diversity is limited at the lowest levels of soil fertility.Location Fragmented, long‐ungrazed Eucalyptus loxophleba subsp. loxophleba (York gum)–Acacia acuminata (jam) woodlands in the wheatbelt of South‐Western Australia.Methods We conducted stratified surveys of floristic composition and topsoil nutrient concentrations in 112 woodland patches. We used generalized linear models, structural equation models and ordinations to characterize relationships among soil nutrients, rainfall, exotic annuals and patch‐scale (100 m2) native plant composition and diversity.Results Patch‐scale native plant diversity declined strongly with increasing exotic abundance. This was partly related to elevated soil nutrient concentrations, particularly total nitrogen and available phosphorus. By contrast, there was little evidence for positive correlations between soil nutrients and native diversity, even at very low soil nutrient concentrations.Main conclusions Minimizing weed invasions is crucial for maximizing native plant diversity in E. loxophleba woodlands and could include nutrient‐depleting treatments without substantially compromising the functional capacity of soils to maintain native plant richness and composition. More broadly we emphasize that understanding relationships among ecosystem productivity, plant diversity and exotic invasions in the context of associated theoretical frameworks is fundamental for informing ecological restoration.

Resource heterogeneity and persistence of exotic annuals in long-ungrazed Mediterranean-climate woodlands

Processes that promote weed invasion are often well-demonstrated, but mechanisms that facilitate ecological resistance to weed invasion in non-invaded communities, or promote weed persistence in invaded communities, are poorly understood. Yet it is these processes that must be addressed to achieve sustainable ecological restoration. We surveyed soil heterogeneity in 25 long-ungrazed, unfertilized York gum (Eucalyptus loxophleba Benth. subsp. loxophleba)—jam (Acacia acuminata Benth.) woodlands of the Western Australian wheatbelt to investigate differences in soil characteristics between patches locally-invaded or non-invaded by widespread exotic annuals. Based on studies in other ecosystems, we hypothesized that (1) weed persistence is associated with elevated soil resource levels, and (2) of these soil resources, phosphorus is the key contributor to weed persistence in Western Australian woodlands, that typically occur on phosphorus-impoverished soils. Our first hypothesis was partly supported, with soil nutrients associated with up to 40% of the variation in cover of exotic annuals. In particular, low concentrations of total nitrogen, nitrate and available phosphorus are likely to contribute to resistance to invasion in many non-invaded woodland patches, especially in gaps between trees. However, other non-invaded patches had comparable nutrient concentrations to invaded patches, suggesting this resistance may be weak at more productive sites or that patches have not reached a stable equilibrium. Inconsistent with our second hypothesis, exotic annuals were as strongly correlated with elevated total nitrogen and associated variables as they were with available phosphorus, probably reflecting a history of grazing without fertilization. We conclude that effectiveness of ‘bottom-up’ approaches to weed control is likely to differ among ecosystems according to interactions with disturbance history and attributes of the non-invaded community, even where the weed species or functional types are the same.