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.     

Restoration of Themeda australis swards suppresses soil nitrate and enhances ecological resistance to invasion by exotic annuals

Understanding processes that underlie ecological resistance to weed invasion is critical for sustainable restoration of invaded plant communities. Experimental studies have demonstrated that invasive nitrophilic annuals can be controlled by addition of carbon to reduce soil nitrate concentrations, sometimes leading to enhanced establishment of native plants. However, effects of carbon supplements on soil nitrate are temporary, and the longer-term value of carbon supplementation as a restoration tool is dependent on the resistance of the re-established ecosystem to repeat invasion. We investigated whether re-established swards of the tussock grass Themeda australis (R.Br.) Stapf (a natural understorey dominant in mesic grassy woodlands of SE Australia) could suppress soil nitrate concentrations, and through this or other means, could impart ongoing resistance to exotic invasion in restored woodlands. In a remnant invaded by exotic annuals, we applied three plot treatments (carbon supplements, annual spring burns and untreated control) and two seed treatments (± Themeda seed) in a replicated, factorial design. Within 3 years, successful establishment of Themeda swards on burnt and carbon-supplemented plots was associated with a reduction in soil nitrate to levels comparable with non-invaded, Themeda-dominated reference sites in the region (<3 mg/kg), and significantly reduced exotic cover compared with unseeded plots. By contrast, on plots not seeded with Themeda, soil nitrate increased after cessation of carbon addition and exotic cover returned to levels comparable with untreated control plots, despite a high cover of other native perennial grasses. Few persistent effects of carbon supplements or spring burning on soil nutrients were evident 9–19 months after cessation of these treatments. Results suggest that Themeda is a keystone species that regulates nitrate cycling, thereby imparting ecological resistance to invasion by nitrophilic annuals.     

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.     

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.     

Invasion resistance and persistence: established plants win,even with disturbance and high propagule pressure

Disturbances and propagule pressure are key mechanisms in plant community resistance to invasion, as well as persistence of invasions. Few studies, however, have experimentally tested the interaction of these two mechanisms. We initiated a study in a southwestern ponderosa pine (Pinus ponderosa Laws.)/bunch grass system to determine the susceptibility of remnant native plant communities to cheatgrass (Bromus tectorum L.) invasion, and persistence of cheatgrass in invaded areas. We used a 2 × 2 factorial design consisting of two levels of aboveground biomass removal and two levels of reciprocal seeding. We seeded cheatgrass seeds in native plots and a native seed mixture in cheatgrass plots. Two biomass removal disturbances and sowing seeds over 3 years did not reverse cheatgrass dominance in invaded plots or native grass dominance in non-invaded native plots. Our results suggest that two factors dictated the persistence of the resident communities. First, bottlebrush squirreltail (Elymus elymoides (Raf.) Swezey) was the dominant native herbaceous species on the study site. This species is typically a poor competitor with cheatgrass as a seedling, but is a strong competitor when mature. Second, differences in pretreatment levels of plant-available soil nitrogen and phosphorus may have favored the dominant species in each community. Annual species typically require higher levels of plant-available soil nutrients than perennial plants. This trend was observed in the annual cheatgrass community and perennial native community. Our study shows that established plants and soil properties can buffer the influences of disturbance and elevated propagule pressure on cheatgrass invasion.     

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.     

Effects of phosphorus on growth and competitive interactions of native and introduced species found in White Box woodlands

Abstract White Box (Eucalyptus albens Benth.) woodlands are among Australia's most endangered ecosystems and are threatened by exotic species invasion. There is evidence from other Australian communities that phosphorus enrichment can facilitate invasion, and differential growth of native and exotic species under increased phosphorus is a possible mechanism. Two glasshouse experiments were designed to test the following three questions relating to species responses to phosphorus: (i) do exotic and native species have different patterns of growth along a gradient of increasing phosphorus?; (ii) do exotic species have a greater competitive effect on native species than do conspecifics?; (iii) does phosphorus enrichment compound the competitive effect of exotic species on native species? Four native perennial species (Themeda australis (R. Br.) Staph., Bothriochloa macra (Steud.) S. T. Blake, Austrodanthonia racemosa (R. Br.) H. P. Linder and Eucalyptus albens) and two exotic annual species (Vulpia bromoides (L) Gray and Echium plantagineum L) were used. In the first experiment, plants were grown individually under six levels of soil phosphorus ranging from 0 to 60 p.p.m. In the second experiment, individuals of Eucalyptus albens and B. macra were grown alone, with a conspecific competitor, or with an exotic (V. bromoides or Echium plantagineum) competitor under low (10 p.p.m.) and high (100 p.p.m.) phosphorus. Both exotic species showed a greater positive response to increased phosphorus than the native species in experiment 1, and Eucalyptus albens seedlings grown with Echium plantagineum were significantly smaller than individuals grown alone or with Eucalyptus albens in experiment 2. There was no evidence that high phosphorus increased the competitive effect of the exotic species, but the combination of a strong positive response to phosphorus and a strong effect on growth of a native species indicates that phosphorus enrichment could favour exotic species in woodland remnants and that field studies testing the effect of phosphorus in a broader context would be appropriate.     

Impact of Parthenium hysterophorus L. invasion on plant species composition and soil properties of grassland communities in Nepal

Parthenium hysterophorus (Asteraceae) is a noxious plant that is considered one of the most invasive species in the world. We studied changes in the composition of plant species and soil properties related to the invasion of P. hysterophorus in three grassland communities of central Nepal. We collected vegetation and soil data along transects that were established in densely invaded to non-invaded areas within homogenous grassland stands. We found significant differences between invaded, transitional and non-invaded plots in species composition and soil properties. There were fewer species in non-invaded than transitional and invaded plots. By P. hysterophorus invasion both native and non-native species were supported or replaced, respectively. The concentrations of soil nitrogen and organic matter were significantly higher in transitional and invaded plots than in non-invaded plots. Soil pH, phosphorus and potassium were highest in the invaded plots, lowest in the non-invaded and intermediate in the transitional plots. Due to changes in above-ground vegetation and below-ground soil nutrient contents, P. hysterophorus invasion is likely to have an overall negative effect on the functioning of the entire ecosystem. Therefore, management of noxious P. hysterophorus is necessary to prevent future problems.     

Understory Invasion by <Emphasis Type="Italic">Acacia longifolia</Emphasis> Alters the Water Balance and Carbon Gain of a Mediterranean Pine Forest

In water-limited ecosystems, where potential evapotranspiration exceeds precipitation, it is often assumed that plant invasions will not increase total ecosystem water use, because all available water is evaporated or transpired regardless of vegetation type. However, invasion by exotic species, with high water use rates, may potentially alter ecosystem water balance by reducing water available to native species, which may in turn impact carbon assimilation and productivity of co-occurring species. Here, we document the impact of invasion by an understory exotic woody species (Acacia longifolia) in a semi-arid Mediterranean dune pine forest. To quantify the effects of this understory leguminous tree on the water use and carbon fixation rates of Pinus pinaster we compare an invaded and a non-invaded stand. A. longifolia significantly altered forest structure by increasing plant density and leaf area index in the mid-stratum of the invaded forest. A. longifolia contributed significantly to transpiration in the invaded forest (up to 42%) resulting in a slight increase in stand transpiration in the invaded relative to non-invaded forest. More importantly, both water use and carbon assimilation rates of P. pinaster were significantly reduced in the invaded relative to non-invaded stand. Therefore, this study shows that exotic plant invasions can have significant impacts on hydrological and carbon cycling even in water-limited semi-arid ecosystems through a repartitioning of water resources between the native and the invasive species.     

Green Frogs Show Reduced Foraging Success in Habitats Invaded by Japanese knotweed

Habitat loss is causing amphibian population declines worldwide, so there is increased attention to forces that degrade remaining habitats. Terrestrial habitats surrounding wetlands are critical foraging areas for temperate anurans. We investigated plant community changes in two old fields invaded by Japanese knotweed (Fallopia japonica) and the foraging success of Green frogs (Rana clamitans) in invaded and non-invaded portions of those fields. Within each field, vegetation data were recorded in quadrats located along two transects bisecting the invasion fronts. We placed frogs in ‘foraging buckets’ along transects and measured their change in mass over a 38 h period. There were significant changes in vegetation structure and composition associated with Japanese knotweed invasion. Diverse assemblages of native plants that covered non-invaded plots were absent from areas invaded by Japanese knotweed. There was also a significant change in vegetation architecture between invaded and non-invaded habitats. Change in frog mass declined significantly along transects, with most frogs in non-invaded plots gaining mass and no frogs in invaded plots gaining mass. Most frogs from non-invaded plots but only two from invaded plots defecated shortly after removal from foraging buckets (verification of recent feeding). We hypothesize that Japanese knotweed invasions degrade terrestrial habitat quality for frogs by indirectly reducing arthropod abundance. Nonnative plant invasions may be another factor contributing to amphibian population declines.     

Effects of exotic grass invasion on spatial heterogeneity in the ground-layer of grassy woodlands

We quantified the effects of exotic annual grass invasion on the ground-layer structure of grassy eucalypt woodlands, with the aim of determining if weed invasion decreased gap size and plant basal area leading to reduced spatial heterogeneity. We measured plant density, distance between plants and basal plant area in woodland sites which ranged from zero to 100% exotic plant cover in the ground-layer. The ground-layer in uninvaded woodlands was heterogeneous, with a large variation in basal plant area and distance between plants. Exotic annual grass density was positively correlated with total plant density, whereas native plant density was negatively correlated. Total plant basal area decreased as total plant density increased, with a lower total plant area in exotic dominated transects compared to native dominated. Variation in basal plant area decreased with increasing plant density. Exotic annual grasses were more closely spaced together (smaller gap size) and had a smaller basal area than the native grasses and rushes. There was also less variation in basal area and gap size with individual exotic annual grasses compared to the native grasses. Inter-plant distance was greater for both the native and exotic grasses when they had native grasses neighbouring them instead of exotic grasses. These findings show that woodlands invaded by exotic annual grasses have relatively less spatial heterogeneity in the ground-layer. These results have implications for other aspects of perennial grassy ecosystems invaded by annual grasses, including plant recruitment and restoration strategies.     

An invasive aster (Ageratina adenophora) invades and dominates forest understories in China: altered soil microbial communities facilitate the invader and inhibit natives

Exotic plant invasion may alter underground microbial communities, and invasion-induced changes of soil biota may also affect the interaction between invasive plants and resident native species. Increasing evidence suggests that feedback of soil biota to invasive and native plants leads to successful exotic plant invasion. To examine this possible underlying invasion mechanism, soil microbial communities were studied where Ageratina adenophora was invading a native forest community. The plant–soil biota feedback experiments were designed to assess the effect of invasion-induced changes of soil biota on plant growth, and interactions between A. adenophora and three native plant species. Soil analysis showed that nitrate nitrogen (NO₃⁻-N), ammonium nitrogen (NH₄⁺-N), and available P and K content were significantly higher in a heavily invaded site than in a newly invaded site. The structure of the soil microbial community was clearly different in all four sites. Ageratina adenophora invasion strongly increased the abundance of soil VAM (vesicular-arbuscular mycorrhizal fungi) and the fungi/bacteria ratio. A greenhouse experiment indicated that the soil biota in the heavily invaded site had a greater inhibitory effect on native plant species than on A. adenophora and that soil biota in the native plant site inhibited the growth of native plant species, but not of A. adenophora. Soil biota in all four sites increased A. adenophora relative dominance compared with each of the three native plant species and soil biota in the heavily invaded site had greater beneficial effects on A. adenophora relative dominance index (20% higher on average) than soil biota in the non-invaded site. Our results suggest that A. adenophora is more positively affected by the soil community associated with native communities than are resident natives, and once the invader becomes established it further alters the soil community in a way that favors itself and inhibits natives, helping to promote the invasion. Soil biota alteration after A. adenophora establishment may be an important part of its invasion process to facilitate itself and inhibit native plants.     

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.     

Impacts of a woody invader vary in different vegetation communities

The impact of an exotic species in natural systems may be dependent not only on invader attributes but also on characteristics of the invaded community. We examined impacts of the invader bitou bush, Chrysanthemoides monilifera ssp. rotundata , in fore and hind dune communities of coastal New South Wales, Australia. We compared invader impacts on vegetation structure, richness of both native and exotic growth forms and community variability in fore and hind dunes. We found that impacts of bitou invasion were context specific: in fore dune shrublands, functionally distinct graminoid, herb and climber rather than shrub growth forms had significantly reduced species richness following bitou invasion. However, in forested hind dunes, the functionally similar native shrub growth form had significantly reduced species richness following bitou invasion. Density of vegetation structure increased at the shrub level in both fore and hind dune invaded communities compared with non-invaded communities. Fore dune ground-level vegetation density declined at invaded sites compared with non-invaded sites, reflecting significant reductions in herb and graminoid species richness. Hind dune canopy-level vegetation density was reduced at invaded compared with non-invaded sites. Bitou bush invasion also affected fore dune community variability with significant increases in variability of species abundances observed in invaded compared with non-invaded sites. In contrast, variability among all hind dune sites was similar. The results suggest that effects of bitou bush invasion are mediated by the vegetation community. When bitou bush becomes abundant, community structure and functioning may be compromised.     

Soil biota composition and the performance of a noxious weed across its invaded range

The success of invasive plant species is driven, in part, by feedback with soil ecosystems. Yet, how variation in belowground communities across latitudinal gradients affects invader distributions remains poorly understood. To determine the effect of soil communities on the performance of the noxious weed Cirsium arvense across its invaded range, we grew seedlings for 40 days in soils collected across a 699 km linear distance from both inside and outside established populations. We also described the mesofaunal and bacterial communities across all soil samples. We found that C. arvense typically performed better when grown in soils sourced from northern populations than from southern locations where it has a longer invasion history. We also found evidence that C. arvense performed best in soils sourced from outside invaded patches, although this was not consistent across all sites. The bacterial community showed a significant increase in the magnitude of compositional change in invaded sites at higher latitudes, while the mesofaunal community showed the opposite pattern. Bacterial community composition was significantly correlated with C. arvense performance, although mesofaunal community composition was not. Our results demonstrate that the interactions between an invasive plant and associated soil communities change across the invaded range, and the bacterial community in particular may affect variation in plant performance. Observed patterns may be caused by C.arvense presence and time since invasion allowing for an accumulation of species‐specific pathogens in southern soils, while the naïveté of northern soils to invasion results in a more responsive bacterial community. Although these interactions are difficult to predict, such effects could possibly facilitate the establishment of this exotic species to novel locations.     

The Effect of a Disturbance Corridor on an Ecological Reserve

The effect of a pipeline corridor constructed through an ecological reserve in Southern California was investigated by assessing plant species composition and soil chemistry. A homogeneous plant community comprised primarily of exotic annuals was found along the entire length of the corridor. This community has low similarity to the adjacent native plant communities. Soil organic matter was significantly less on the disturbed corridor than in contiguous undisturbed areas. Both available nitrogen and extractable phosphorus values were greater in the disturbed corridor. By contrast, total nitrogen was significantly higher outside the pipeline. The more labile litter of the exotic annuals allows increased mineralization along the corridor than does the more recalcitrant litter of the native perennial shrubs in the undisturbed areas. Once established, the weedy exotic annual litter may completely turn over organic matter and nitrogen, favoring the persistence of the weedy annuals. These exotic annuals appear to be moving into three of the native communities - grassland, coastal sage, and oak woodland - that have less organic matter and a more open plant canopy. Poor restoration efforts can lead to the establishment of such exotics, subsequent invasion into the surrounding undisturbed habitat, and degradation of the reserve.     

Spatial and Temporal Dynamics of Exotic Earthworm Communities Along Invasion Fronts in a Temperate Hardwood Forest in South-Central New York (USA)

The invasion of North American forests by exotic earthworms is producing profound ecosystem changes, such as alterations in soil nutrient cycling, and redistribution and loss of soil organic matter. However, the present and future extent of these invasions is difficult to evaluate without a better understanding of the factors that control the distribution and abundance of earthworms in previously non-invaded habitats. In this study, the species composition and short-term dynamics of three exotic earthworm invasion fronts were studied at a northern hardwood forest in south-central New York State (USA). Belt transects were established at each of the three locations to sample from earthworm-invaded areas through transition zones and into invasion front areas. Lumbricus rubellus, L. terrestrisandOctolasion tyrtaeum were the most common species, but their distribution was not homogeneous along the transects. Whereas, L. rubellus was the only species with relatively high adult densities at transition zones and invasion fronts, L. terrestris and O. tyrtaeum occurred mostly in the heavily earthworm-invaded areas and were rare at the invasion fronts. The density of earthworms along the transects decreased by 60–87 from June 2001 to October 2002 and then recovered in 2003 to values similar to those of 2001. This decrease was apparently caused by reduced recruitment of immature earthworms, probably related to the severe drought periods that the study area experienced in 2001 and 2002. Our data suggest that climate and topography, through their effects on soil moisture patterns, can be critical factors controlling the distribution and spread of exotic earthworms in previously non-invaded habitats.     

Identification of volatile compounds released by roots of an invasive plant, bitou bush (Chrysanthemoides monilifera spp. rotundata), and their inhibition of native seedling growth

Allelopathy has been suggested as a mechanism promoting the monoculture formation of some invasive exotic plants. Previous studies have shown that hydrophobic extracts of the roots and soil of exotic bitou bush (Chrysanthemoides monilifera spp. rotundata (DC.) T. Norl.) inhibited the seedling growth of five Australian native plants, including the dominant acacia (Acacia longifolia var. sophorae (Labill.) F. Muell.). Based on this finding, we compared the hydrophobic root and soil chemical profiles of bitou bush and acacia to determine whether bitou bush roots release allelopathic compounds that are novel to the invaded system. We detected three compounds that were exclusive to the bitou bush root and soil, and seven compounds that were common to the bitou bush and acacia roots but only present in the bitou bush soil. The compounds unique to the bitou bush invaded soil were all sesqui- and diterpenes. Several of these compounds were found to inhibit the seedling growth of a native sedge, Isolepis nodosa (Rott.) R. Br. Of particular interest are the sesquiterpenes: β-maaliene, α-isocomene, β-isocomene, δ-cadinene, 5-hydroxycalamenene and 5-methoxycalamenene which were found in high concentrations in the bitou bush root and soil extracts and exhibited phytotoxic activity. Therefore, we present evidence to suggest that bitou bush exudes low molecular weight volatile compounds into the soil which inhibit native plant seedling growth. The reduced establishment of native plants via allelopathy is likely to create space and contribute to the invasion of bitou bush on the eastern Australian coast.     

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.     

Seed predation of two alien Opuntia species invading Mediterranean communities

Invasibility depends on the interaction of the introduced species with the abiotic and biotic factors of the recipient community. In particular, the biotic resistance posed by native herbivores has been claimed to be of great importance in controlling plant invasion. We investigated fruit and seed predation of two exotic Opuntia species within and between Mediterranean communities in order to determine how patterns of predation matched patterns of invasion. Predators were small mammals, presumably mice, which could consume more than 50% of the seeds produced. Predators could be equally effective in consuming fruit and single seeds. O. maxima fruits were slightly preferred to O. stricta fruits, but predators did not distinguish between seeds. Seed predation was more intense in invaded than in non-invaded communities. However, there was a high spatial variation in seed predation that did not always match patterns of invasion, suggesting that seed predation alone is not a good predictor of community invasibility to Opuntia. According to these results invasibility to Opuntia is limited in some (but not all) communities by native mice. Seed losses by predation were high for both species. However, we estimated that more than 75% of seeds dispersed by birds to non-invaded areas are not predated.