Establishment of native grasses and their impact on exotic annuals in degraded box gum woodlands

Restoration goals often involve the addition of new species to resident, degraded communities but in box gum woodlands such restoration is often constrained by competition from persistent exotic annuals that control critical ecological processes. Nutrient reduction (via carbon addition) and seed bank depletion are two approaches to reduce competition from exotic annuals but to be effective these treatments must allow establishment of species such as native grasses. This experiment was conducted in two degraded Austrostipa understoreys in the box gum woodlands of south‐east Australia. It compares the effects of carbon addition (sugar), seed depletion (spring burning or spring grazing) and combinations of carbon addition and seed depletion treatments on the establishment of C3 and C4 native grasses, and measured the effects of their establishment on soil nitrate concentration and exotic annuals. Treatments that reduced exotic annual abundance did not increase initial germination of the C4 native grasses, Bothriochloa or Themeda. However, sugar increased seedling survival of Themeda and Bothriochloa and grazing increased seedling survival of Bothriochloa, presumably by reducing effects of exotic annuals. Poa and Rytidosperma (C3 native grasses) failed to establish. Although we were unable to detect any reduction in soil nitrate concentration, swards with successful recruitment of C4 grasses suppressed exotic annuals more than the Austrostipa‐only swards at one site (the other was affected by wildfire). Further, Austrostipa‐Themeda swards were more effective than Austrostipa‐Bothriochloa for suppressing exotics, pointing to a role for both functional and species identity in the degree of resistance conferred.     

A plant traits approach to managing legacy species during restoration transitions in temperate eucalypt woodlands

Degraded communities often contain a subset of the species that comprised the predisturbance community. These represent an important legacy of the predisturbance state, yet restoration treatments may be detrimental to them. This study examined the potential of leaf traits and life form to predict whether restoration treatments can maintain legacy swards of Austrostipa bigeniculata (hereafter Austrostipa) while controlling exotic annuals in temperate eucalypt woodlands. Treatments included carbon addition to reduce soil nitrate, both with and without burning or pulse grazing to deplete exotic seed pools. We compared leaf traits of Austrostipa with a native grass (Themeda triandra) known to be advantaged, and 8 exotic annual species known to be disadvantaged by these treatments. Leaf traits indicated potentially greater negative impacts of carbon addition on exotic annuals compared to Austrostipa, and on Austrostipa compared to Themeda, suggesting a net restoration benefit. Similarly, burning or pulse grazing is expected to have little negative impact on perennial resprouting grasses (hemicryptophytes; Austrostipa and Themeda) compared with annual exotics (therophytes) with short‐lived seed banks. Treatment responses were largely consistent with predictions: treatments that significantly reduced exotic annuals had no net disadvantage to Austrostipa swards despite significant reductions in Austrostipa seedling growth with carbon addition. Indeed by Year 3, Austrostipa mortality in untreated plots led to 46% lower Austrostipa abundance than in treated plots at one site, potentially due to litter build‐up or other mechanisms. We conclude that plant traits provide a useful framework for designing restoration transitions that retain native legacy species while controlling exotics.     

Effects of Sowing Treatment and Landscape Position on Establishment of the Perennial Tussock Grass Themeda triandra (Poaceae) in Degraded Eucalyptus Woodlands in Southeastern Australia

Grassy woodlands dominated by Eucalyptus species such as E. albens Benth., E. melliodora A. Cunn. ex Schauer, and E. microcarpa Maiden are poorly conserved in Australia, owing to widespread agricultural development. Understorey restoration is vitally important to enhance degraded remnants but no reliable techniques are available to restore herbaceous understorey species over large areas. Reestablishment of dominant native grasses such as Kangaroo grass (Themeda triandra Forssk.) is particularly important. This study compared Themeda establishment using a variety of sowing techniques across a range of landscape positions in degraded woodlands in central New South Wales. Four localities were sampled at three landscape positions (upper, mid‐, and lower slopes). Existing vegetative cover was sprayed with herbicide and removed by mowing. A randomized block experiment was established at each site with five replicates of five seedbed treatments: all four factorial combinations of soil disturbance (disturbed, nondisturbed) and weed control (atrazine, nonatrazine), plus topsoil scalping followed by soil disturbance. After 117 days, all plots were subject to a common herbicide (atrazine) treatment to control weeds. Themeda establishment was not significantly affected by landscape position, despite significant differences in the cover of exotic species among landscape positions. The scalped/disturbed treatment resulted in significantly better establishment at 400 days (18% or 3.9 plants/m²) than the control (8% or 1.8 plants/m²). All other treatments did not differ significantly from the control suggesting that seedbed treatments are not critical to successful Themeda establishment. These results suggest that Themeda swards can be reestablished in degraded woodlands relatively simply. The similarity in outcomes across all landscape positions suggests that general recommendations for “best‐bet” establishment methods may prove robust under a wide range of environmental and seasonal conditions.     

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.     

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

Aim 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.     

Giant invasive Heracleum persicum: Friend or foe of plant diversity?

The impact of invasion on diversity varies widely and remains elusive. Despite the considerable attempts to understand mechanisms of biological invasion, it is largely unknown whether some communities’ characteristics promote biological invasion, or whether some inherent characteristics of invaders enable them to invade other communities. Our aims were to assess the impact of one of the massive plant invaders of Scandinavia on vascular plant species diversity, disentangle attributes of invasible and noninvasible communities, and evaluate the relationship between invasibility and genetic diversity of a dominant invader. We studied 56 pairs of Heracleum persicum Desf. ex Fisch.‐invaded and noninvaded plots from 12 locations in northern Norway. There was lower native cover, evenness, taxonomic diversity, native biomass, and species richness in the invaded plots than in the noninvaded plots. The invaded plots had nearly two native species fewer than the noninvaded plots on average. Within the invaded plots, cover of H. persicum had a strong negative effect on the native cover, evenness, and native biomass, and a positive association with the height of the native plants. Plant communities containing only native species appeared more invasible than those that included exotic species, particularly H. persicum. Genetic diversity of H. persicum was positively correlated with invasibility but not with community diversity. The invasion of a plant community by H. persicum exerts consistent negative pressure on vascular plant diversity. The lack of positive correlation between impacts and genetic diversity of H. persicum indicates that even a small founder population may cause high impact. We highlight community stability or saturation as an important determinant of invasibility. While the invasion by H. persicum may decrease susceptibility of a plant community to further invasion, it severely reduces the abundance of native species and makes them more vulnerable to competitive exclusion.     

Effectiveness of repeated autumn and spring fires for understorey restoration in weed‐invaded temperate eucalypt woodlands

Question. Can strategic burning, targeting differing ecological characteristics of native and exotic species, facilitate restoration of native understorey in weed‐invaded temperate grassy eucalypt woodlands? Location. Gippsland Plains, eastern Victoria, Australia. Methods. In a replicated, 5‐year experimental trial, the effects of repeated spring or autumn burning were evaluated for native and exotic plants in a representative, degraded Eucalyptus tereticornis grassy woodland. Treatments aimed to reduce seed banks and modify establishment conditions of exotic annual grasses, and to exhaust vegetative reserves of exotic perennial grasses. Treatments were applied to three grassland patch types, dominated by the native grass Austrodanthonia caespitosa, ubiquitous exotic annuals, or the common exotic perennial grass Paspalum dilatatum. Results. The dominant native grass Austrodanthonia caespitosa and native forbs were resilient to repeated fires, and target exotic annuals and perennials were suppressed differentially by autumn and spring fires. Exotic annuals were also suppressed by drought, reducing the overall treatment effects but indicating important opportunities for restoration. The initially sparse exotic geophyte Romulea rosea increased in cover with fire and the impact of this species on native forbs requires further investigation. There was minimal increase in diversity of subsidiary natives with fire, probably owing to lack of propagules. Conclusions. While fire is often considered to increase ecosystem invasibility, our study showed that strategic use of fire, informed by the relative responses of available native and exotic taxa, is potentially an effective step towards restoration of weed‐invaded temperate eucalypt woodlands.     

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

Control 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.     

Patterns of Plant Invasions: A Case Example in Native Species Hotspots and Rare Habitats

Land managers require landscape-scale information on where exotic plant species have successfully established, to better guide research, control, and restoration efforts. We evaluated the vulnerability of various habitats to invasion by exotic plant species in a 100,000 ha area in the southeast corner of Grand Staircase-Escalante National Monument, Utah. For the 97 0.1-ha plots in 11 vegetation types, exotic species richness (log₁₀) was strongly negatively correlated to the cover of cryptobiotic soil crusts (r = −0.47, P < 0.001), and positively correlated to native species richness (r = 0.22, P < 0.03), native species cover (r = 0.23, P < 0.05), and total nitrogen in the soil (r = 0.40, P < 0.001). Exotic species cover was strongly positively correlated to exotic species richness (r = 0.68, P < 0.001). Only 6 of 97 plots did not contain at least one exotic species. Exotic species richness was particularly high in locally rare, mesic vegetation types and nitrogen rich soils. Dry, upland plots (n = 51) had less than half of the exotic species richness and cover compared to plots (n = 45) in washes and lowland depressions that collect water intermittently. Plots dominated by trees had significantly greater native and exotic species richness compared to plots dominated by shrubs. For the 97 plots combined, 33% of the variance in exotic species richness could be explained by a positive relationship with total plant cover, and negative relationships with the cover of cryptobiotic crusts and bare ground. There are several reasons for concern: (1) Exotic plant species are invading hot spots of native plant diversity and rare/unique habitats. (2) The foliar cover of exotic species was greatest in habitats that had been invaded by several exotic species.(3) Continued disturbance of fragile cryptobiotic crusts by livestock, people, and vehicles may facilitate the further invasion of exotic plant species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evaluating strategies for facilitating native plant establishment in northern Nevada crested wheatgrass seedings

Non‐native crested wheatgrasses (Agropyron cristatum and A. desertorum) were used historically within the Great Basin for the purpose of competing with weed species and increasing livestock forage. These species continue to be used in some areas, especially after wildfires occurring in low elevation/precipitation, formerly Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis)/herbaceous communities. Seeding native species in these sites is often unsuccessful, and lack of establishment results in invasion and site dominance by exotic annuals. However, crested wheatgrass often forms dense monocultures that interfere competitively with the establishment of desirable native vegetation and do not provide the plant structure and habitat diversity for wildlife species equivalent to native‐dominated sagebrush plant communities. During a 5‐year study, we conducted trials to evaluate chemical and mechanical methods for reducing crested wheatgrass and the effectiveness of seeding native species into these sites after crested wheatgrass suppression. We determined that discing treatments were ineffective in reducing crested wheatgrass cover and even increased crested wheatgrass density in some cases. Glyphosate treatments initially reduced crested wheatgrass cover, but weeds increased in many treated plots and seeded species diminished over time as crested wheatgrass recovered. We concluded that, although increases in native species could possibly be obtained by repeating crested wheatgrass control treatments, reducing crested wheatgrass opens a window for invasion by exotic weed species.     

The Effects of Organic Amendments on the Restoration of a Disturbed Coastal Sage Scrub Habitat

The effectiveness of organic mulch as a simple means of enhancing the restoration of disturbed lands by providing a competitive edge to native perennials, such as Artemisia californica (California sagebrush), over exotic annuals, such as Avena fatua (wild oat), was studied by investigating the effect of organic amendments on microbial activity and nitrogen immobilization through both soil analysis and aboveground plant growth. The addition of organic amendment resulted in an increase in microbial activity, a parallel increase in nitrogen immobilization, and no significant differences in total soil nitrogen. It is likely that nitrogen was gradually being removed from its more available form of nitrate and being immobilized in the tissues of the increasing microbial biomass. The survival rate of planted native perennial seedlings of A. californica in organic amended plots was almost double that of control-plot seedlings, and plant volume was significantly higher. When the availability of nitrogen was reduced through increased immobilization, amended plots established an environment more conducive to native perennial shrubs, allowing them to outcompete exotic annuals for water and nutrients. This simple procedure could have major implications for enhancing the restoration of disturbed lands.     

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

Shrub 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 & 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.     

Net carbon exchange and evapotranspiration in postfire and intact sagebrush communities in the Great Basin

Invasion of non-native annuals across the Intermountain West is causing a widespread transition from perennial sagebrush communities to fire-prone annual herbaceous communities and grasslands. To determine how this invasion affects ecosystem function, carbon and water fluxes were quantified in three, paired sagebrush and adjacent postfire communities in the northern Great Basin using a 1-m³ gas exchange chamber. Most of the plant cover in the postfire communities was invasive species including Bromus tectorum L., Agropyron cristatum (L.) Gaertn and Sisymbrium altissimum L. Instantaneous morning net carbon exchange (NCE) and evapotranspiration (ET) in native shrub plots were greater than either intershrub or postfire plots. Native sagebrush communities were net carbon sinks (mean NCE 0.2–4.3 μmol m⁻² s⁻¹) throughout the growing season. The magnitude and seasonal variation of NCE in the postfire communities were controlled by the dominant species and availability of soil moisture. Net C exchange in postfire communities dominated by perennial bunchgrasses was similar to sagebrush. However, communities dominated by annuals (cheatgrass and mustard) had significantly lower NCE than sagebrush and became net sources of carbon to the atmosphere (NCE declined to −0.5 μmol m⁻² s⁻¹) with increased severity of the summer drought. Differences in the patterns of ET led to lower surface soil moisture content and increased soil temperatures during summer in the cheatgrass-dominated community compared to the adjacent sagebrush community. Intensive measurements at one site revealed that temporal and spatial patterns of NCE and ET were correlated most closely with changes in leaf area in each community. By altering the patterns of carbon and water exchange, conversion of native sagebrush to postfire invasive communities may disrupt surface-atmosphere exchange and degrade the carbon storage capacity of these systems.     

Restoring a Mediterranean‐climate shrub community with perennial species reduces future invasion

Successful restoration of an invaded landscape to a diverse, invasion‐resistant native plant community requires determining the optimal native species mix to add to the landscape. We manipulated native seed mix (annuals, perennials, or a combination of the two), while controlling the growth of non‐native species to test the hypothesis that altering native species composition can influence native establishment and subsequent non‐native invasion. Initial survival of native annuals and perennials was higher when seeded in separate mixes than when combined, and competition between the native perennials and annuals led to lower perennial cover in year 2 of mixed‐seeded plots. The plots with the highest perennial cover had the highest resistance to invasion by Brassica nigra. To clarify interactions among different functional groups of natives and B. nigra, we measured competitive interactions in pots. We grew one native annual, one native perennial, and B. nigra alone or with different competitors and measured biomass after 12 weeks. Brassica nigra was the strongest competitor, limiting the growth of all native species, and was not impacted by competition with native annuals or perennial seedlings. Results from the potted plant experiment demonstrated the strong negative influence of B. nigra on native seedlings. Older native perennials were the strongest competitors against invasive species in the field, yet perennial seedling survival was limited by competition with native annuals and B. nigra. Management action that maximizes perennial growth in early years may lead to a relatively more successful restoration and the establishment of an invasion‐resistant community.     

Invasion by an exotic tree alters above and belowground ecosystem components

With the widespread introduction and invasion of exotic plants there is a need for studies that quantify alterations of basic ecosystem structure and function. Ecosystem invasion by Melaleuca quinquenervia significantly altered both above- and belowground ecosystem components in this study. We measured the quantity and nutrient concentration of the litterfall, litter layer, and soil; microbial biomass pools; and rates of potentially mineralizable nitrogen and soil oxygen demand. Annual litterfall was 4.9 times higher in the non-invaded sites and contained 1.9 times more phosphorus than invaded sites. Non-invaded plots contained a larger litter layer compared to invaded plots: 2.4 ± 1.2 kg m⁻² and 0.62 ± 0.3 kg m⁻² , respectively. Lower nutrient concentration and quantity of the litter layer in the invaded plots led to changes in the aboveground storage of nutrients. In the invaded plots there was four times less carbon, seven times less nitrogen, and ten times less phosphorus stored in the organic litter layer compared to the non-invaded plots. Microbial biomass nutrient pools were consistently lower at both the 0–5 cm and 5–15 cm depth in the invaded soils compared to non-invaded soils, indicating a plant mediated change. Although M. quinquenervia altered microbial community structure, microbial activities were not different between invaded and non-invaded plots at either depth as measured by rates of soil oxygen demand and potentially mineralizable nitrogen. These changes may affect both native plant growth and water quality, and may act to promote and maintain site dominance by M. quinquenervia.     

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

One 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.     

Riparian zones as havens for exotic plant species in the central grasslands

In the Central Grasslands of the United States, we hypothesized that riparian zones high in soil fertility would contain more exotic plant species than upland areas of low soil fertility. Our alternate hypothesis was that riparian zones high in native plant species richness and cover would monopolize available resources and resist invasion by exotic species. We gathered nested-scale vegetation data from 40 1 m²subplots (nested in four 1000 m² plots) in both riparian and upland sites at four study areas in Colorado, Wyoming, and South Dakota (a total of 320 1 m² subplots and 32 1000 m² plots). At the 1 m² scale, mean foliar cover of native species was significantly greater (P<0.001) in riparian zones (36.3% ± 1.7%) compared to upland sites (28.7% ± 1.5%), but at this small scale there were no consistent patterns of native and exotic species richness among the four management areas. Mean exotic species cover was slightly higher in upland sites compared to riparian sites (9.0% ± 3.8% versus 8.2% ± 3.0% cover). However, mean exotic species richness and cover were greater in the riparian zones than upland sites in three of four management areas. At the 1000 m² scale, mean exotic species richness was also significantly greater (P<0.05) in riparian zones (7.8 ± 1.0 species) compared to upland sites (4.8 ± 1.0 species) despite the heavy invasion of one upland site. For all 32 plots combined, 21% of the variance in exotic species richness was explained by positive relationships with soil % silt (t =1.7, P=0.09) and total foliar cover (t = 2.4, P=0.02). Likewise, 26% of the variance in exotic species cover (log₁₀ cover) was explained by positive relationships with soil % silt (t =2.3, P=0.03) and total plant species richness (t = 2.5, P=0.02). At landscape scales (four 1000 m² plots per type combined), total foliar cover was significantly and positively correlated with exotic species richness (r=0.73, P<0.05) and cover (r=0.74, P<0.05). Exotic species cover (log₁₀ cover) was positively correlated with log₁₀% N in the soil (r=0.61, P=0.11) at landscape scales. On average, we found that 85% (±5%) of the total number of exotic species in the sampling plots of a given management area could be found in riparian zones, while only 50% (±8%) were found in upland plots. We conclude that: (1) species-rich and productive riparian zones are particularly invasible in grassland ecosystems; and (2) riparian zones may act as havens, corridors, and sources of exotic plant invasions for upland sites and pose a significant challenge to land managers and conservation biologists.     

From protégé to nurse plant: establishment of thorny shrubs in grazed temperate woodlands

Question: Thorny shrubs play keystone roles in grazed ecosystems by defending non‐protected plants against herbivores, but their establishment in grazed ecosystems is poorly understood. Which factors control establishment of recruits of thorny nurse shrubs in grazed temperate woodlands? Location: Ancient grazed temperate woodlands (52°32′N, 6°36′E), The Netherlands. Methods: We surveyed biotic and abiotic factors for saplings of thorny nurse shrubs in plots with and without saplings. To disentangle these factors, we performed a transplantation experiment over two growing seasons with nurse shrub saplings (Prunus spinosa and Crataegus monogyna) planted in two dominant vegetation types – tall unpalatable swards and short grazed lawns – half of them protected from herbivory via exclosures. Results: Plots with shrub saplings had taller surrounding vegetation, higher soil pH and higher soil moisture than plots without saplings. These plots predominantly contained unpalatable sward species, while plots without saplings mainly contained palatable lawn species. After transplantation, sapling survival was higher in exclosures than in the open, and higher in sward exclosures than in lawn exclosures. Sapling growth was higher in swards than in lawns, higher inside than outside exclosures, and higher for Prunus than Crataegus, while browsing on saplings was higher in lawns. Conclusion: Unpalatable swards form essential establishment niches for thorny shrubs in grazed temperate woodlands: they protect against herbivores before thorns fully develop in saplings, and sapling growth is better due to improved micro‐environmental conditions. Once established and thorny, shrub saplings grow out of the protective range of the swards and in turn facilitate tree seedlings, which are essential for long‐term persistence of grazed temperate woodlands. This study shows that nurse plants may start as protégés before becoming facilitators for other plants in a later life stage. This may be common for nurse plants in various ecosystems. We argue that improved understanding of establishment of nurse plants and their constraining factors is crucial for effective conservation and restoration in various ecosystems.     

Shrub canopies influence soil temperatures but not nutrient dynamics: An experimental test of tundra snow–shrub interactions

Shrubs are the largest plant life form in tundra ecosystems; therefore, any changes in the abundance of shrubs will feedback to influence biodiversity, ecosystem function, and climate. The snow–shrub hypothesis asserts that shrub canopies trap snow and insulate soils in winter, increasing the rates of nutrient cycling to create a positive feedback to shrub expansion. However, previous work has not been able to separate the abiotic from the biotic influences of shrub canopies. We conducted a 3‐year factorial experiment to determine the influences of canopies on soil temperatures and nutrient cycling parameters by removing ~0.5 m high willow (Salix spp.) and birch (Betula glandulosa) shrubs, creating artificial shrub canopies and comparing these manipulations to nearby open tundra and shrub patches. Soil temperatures were 4–5°C warmer in January, and 2°C cooler in July under shrub cover. Natural shrub plots had 14–33 cm more snow in January than adjacent open tundra plots. Snow cover and soil temperatures were similar in the manipulated plots when compared with the respective unmanipulated treatments, indicating that shrub canopy cover was a dominant factor influencing the soil thermal regime. Conversely, we found no strong evidence of increased soil decomposition, CO₂ fluxes, or nitrate or ammonia adsorbtion under artificial shrub canopy treatments when compared with unmanipulated open tundra. Our results suggest that the abiotic influences of shrub canopy cover alone on nutrient dynamics are weaker than previously asserted.     

Responses of native and non-native Mojave Desert winter annuals to soil disturbance and water additions

Habitat modification (i.e., disturbance) and resource availability have been identified as possible mechanisms that may influence the invasibility of plant communities. In the Mojave Desert, habitat disturbance has increased dramatically over the last 50 years due to increased human activities. Additionally, water availability is considered to be a main limiting resource for plant production. To elucidate the effects of soil disturbance and water availability on plant invasions, we created experimental patches where we varied the levels of soil disturbance and water availability in a fully crossed factorial experiment at five replicated field sites, and documented responses of native and non-native winter annuals. The treatments did not significantly affect the density (seedlings m⁻²) of the non-native forb, Brassica tournefortii. However, the relationship between silique production and plant height differed among treatments, with greater silique production in disturbed plots. In contrast to Brassica, density of the non-native Schismus spp. increased in soil disturbed and watered plots, and was greatest in disturbed plots during 2009 (the second year of the study). Species composition of the native annual community was not affected by treatments in 2008 but was influenced by treatments in 2009. The native forb Eriophyllum sp. was most dense on water-addition plots, while density of Chaenactis freemontii was highest in disturbed plots. Results illustrate that habitat invasibility in arid systems can be influenced by dynamics in disturbance regimes and water availability, and suggest that invasiveness can differ between non-native annual species and among native annuals in habitats undergoing changing disturbance and precipitation regimes. Understanding the mechanistic relationships between water availability and non-native plant responses will be important for understanding the effects of shifting precipitation and vegetation patterns under predicted climate change in arid ecosystems.