Influence of fire and soil nutrients on native and non-native annuals at remnant vegetation edges in the Western Australian wheatbelt

Abstract. The effect of fire on annual plants was examined in two vegetation types at remnant vegetation edges in the Western Australian wheatbelt. Density and cover of non-native species were consistently greatest at the reserve edges, decreasing rapidly with increasing distance from reserve edge. Numbers of native species showed little effect of distance from reserve edge. Fire had no apparent effect on abundance of non-natives in Allocasuarina shrubland but abundance of native plants increased. Density of both non-native and native plants in Acacia acuminata-Eucalyptus loxophleba woodland decreased after fire. Fewer non-native species were found in the shrubland than in the woodland in both unburnt and burnt areas, this difference being smallest between burnt areas. Levels of soil phosphorus and nitrate were higher in burnt areas of both communities and ammonium also increased in the shrubland. Levels of soil phosphorus and nitrate were higher at the reserve edge in the unburnt shrubland, but not in the woodland. There was a strong correlation between soil phosphorus levels and abundance of non-native species in the unburnt shrubland, but not after fire or in the woodland. Removal of non-native plants in the burnt shrubland had a strong positive effect on total abundance of native plants, apparently due to increases in growth of smaller, suppressed native plants in response to decreased competition. Two native species showed increased seed production in plots where non-native plants had been removed. There was a general indication that, in the short term, fire does not necessarily increase invasion of these communities by non-native species and could, therefore be a useful management tool in remnant vegetation, providing other disturbances are minimised.     

Non-native plants and soil microbes: potential contributors to the consistent reduction in soil aggregate stability caused by the disturbance of North American grasslands

? Soil aggregate stability is an important ecosystem property that is altered by anthropogenic disturbance. Yet, the generalization of these alterations and the identification of the main contributors are limited by the absence of cross-site comparisons and the application of inconsistent methodologies across regions. ? We assessed aggregate stability in paired remnant and post-disturbance grasslands across California, shortgrass and tallgrass prairies, and in manipulative experiments of plant composition and soil microbial inoculation. ? Grasslands recovering from anthropogenic disturbance consistently had lower aggregate stability than remnants. Across all grasslands, non-native plant diversity was significantly associated with reduced soil aggregate stability. A negative effect of non-native plants on aggregate stability was also observed in a mesocosm experiment comparing native and non-native plants from California grasslands. Moreover, an inoculation study demonstrated that the degradation of the microbial community also contributes to the decline in soil aggregate stability in disturbed grasslands. ? Anthropogenic disturbance consistently reduced water-stable aggregates. The stability of aggregates was reduced by non-native plants and the degradation of the native soil microbial community. This latter effect might contribute to the sustained decline in aggregate stability following anthropogenic disturbance. Further exploration is advocated to understand the generality of these potential mechanisms.     

Composition and seasonal flux of the soil seed bank of species-rich Themeda triandra grasslands in relation to burning history

Abstract. Most species-rich grasslands dominated by Themeda triandra in southeastern Australia have been ungrazed and frequently burnt for decades. The seedling emergence technique was used to determine the size and taxonomic composition of the soil seed bank of five grasslands that had different fire histories (i.e. burnt at 1 yr, 3 yr and > 10 yr intervals) and this was compared to the standing vegetation at each site. A nested sampling design (subplot, plot, site) was used to determine the effect of spatial scale on the patterns observed in both the vegetation and the seed bank. Temporal variation in the seed bank was assessed by repeated soil sampling over a two year period. 61 native and 30 exotic species were recorded in the vegetation. Richness varied more between sites than within sites. Sites were therefore internally homogeneous for species richness. However, no correlation between burning frequency and richness was found. DCA ordination separated the sites into distinct groups, but sites with similar fire history did not necessarily group closely. 60 taxa germinated from the soil seed bank, comprising 32 native and 28 exotic species; 11 species, mostly therophytes, were restricted to the seed bank. The richness of the seed bank was significantly lower than the vegetation at all spatial scales. No correlation between seed bank richness and fire history was found. The seed bank of species-rich grasslands is dominated by a limited number of widespread, highly clumped, annual, native and exotic monocots. Most native hemicryptophytes, and perennials in general, were represented in the soil by a transient seed bank. Only 12 % of study species, all therophytes, were considered to form large, persistent seed banks, the size of which was greater in unburnt grasslands at all times of the year. The distinct floristic patterns observed in the vegetation are less clearly represented in the seed bank. The seed bank represents a floristically distinct (and less variable) component of the vegetation when compared to the standing flora. The size of the long-term soil seed bank suggests that it has little functional importance for many native species and probably contributes little to seedling regeneration processes following disturbance. Altering established fire regimes is likely to only change the composition and small-scale richness of the existing site vegetation and will not re-integrate species previously lost from the vegetation due to past management. It is suggested that the maintenance of vegetation remnants and processes that encompass a range of long-term burning histories will be necessary if the flora is to be conserved in situ. Restoration of degraded grasslands cannot rely on the soil seed bank but rather, will be dependent on the reintroduction of propagules.     

Reduced soil compaction enhances establishment of non-native plant species

Many studies have shown that soil disturbance facilitates establishment of invasive, non-native plant species, and a number of mechanisms have been isolated that contribute to the process. To our knowledge no studies have isolated the role of altered soil compaction, a likely correlate of many types of soil disturbance, in facilitating invasion. To address this, we measured the response of seeded non-native and native plant species to four levels of soil compaction in mesocosms placed in an abandoned agricultural field in the Methow Valley, Washington, USA. Soil compaction levels reflected the range of resistance to penetration (0.1–3.0 kg cm⁻²) measured on disturbed soils throughout the study system prior to the experiment. Percent cover of non-native species, namely Bromus tectorum and Centaurea diffusa, decreased by 34% from the least to the most compacted treatments, whereas percent cover of native species, mostly Pseudoroegneria spicata and Lupinus spp., did not respond to compaction treatments. Experimental results were supported by a survey of soil penetration resistance and percent cover by species in 18 abandoned agricultural fields. Percent cover of B. tectorum was negatively related to soil compaction levels, whereas none of the native species showed any response to soil compaction. These results highlight a potentially important, though overlooked, aspect of soil disturbance that may contribute to subsequent non-native plant establishment.     

Influence of experimental soil disturbances on the diversity of plants in agricultural grasslands

Aims and Methods Disturbance is supposed to play an important role for biodiversity and ecosystem stability as described by the intermediate disturbance hypothesis (IDH), which predicts highest species richness at intermediate levels of disturbances. In this study, we tested the effects of artificial soil disturbances on diversity of annual and perennial vascular plants and bryophytes in a field experiment in 86 agricultural grasslands differing in land use in two regions of Germany. On each grassland, we implemented four treatments: three treatments differing in application time of soil disturbances and one control. One year after experimental disturbance, we recorded vegetation and measured biomass productivity and bare ground. We analysed the disturbance response taking effects of region and land-use-accompanied disturbance regimes into account.Important findings Region and land-use type strongly determined plant species richness. Experimental disturbances had small positive effects on the species richness of annuals, but none on perennials or bryophytes. Bare ground was positively related to species richness of bryophytes. However, exceeding the creation of 12% bare ground further disturbance had a detrimental effect on bryophyte species richness, which corresponds to the IDH. As biomass productivity was unaffected by disturbance our results indicate that the disturbance effect on species richness of annuals was not due to decreased overall productivity, but rather due to short-term lowered inter- and intraspecific competition at the newly created microsites. Generally, our results highlight the importance of soil disturbances for species richness of annual plants and bryophytes in agricultural grasslands. However, most grasslands were disturbed naturally or by land-use practices and our additional experimental soil disturbances only had a small short-term effect. Overall, total plant diversity in grasslands seemed to be more limited by the availability of propagules rather than by suitable microsites for germination. Thus, nature conservation efforts to increase grassland diversity should focus on overcoming propagule limitation, for instance by additional sowing of seeds, while the creation of additional open patches by disturbance might only be appropriate where natural disturbances are scarce.     

Patterns of invasion of an urban remnant of a species-rich grassland in southeastern Australia by non-native plant species

Abstract. The invasion by non-native plant species of an urban remnant of a species-rich Themeda triandra grassland in southeastern Australia was quantified and related to abiotic influences. Richness and cover of non-native species were highest at the edges of the remnant and declined to relatively uniform levels within the remnant. Native species richness and cover were lowest at the edge adjoining a roadside but then showed little relation to distance from edge. Roadside edge quadrats were floristically distinct from most other quadrats when ordinated by Detrended Correspondence Analysis. Soil phosphorus was significantly higher at the roadside edge but did not vary within the remnant itself. All other abiotic factors measured (NH₄, NO₃, S, pH and % organic carbon) showed little variation across the remnant. Non-native species richness and cover were strongly correlated with soil phosphorus levels. Native species were negatively correlated with soil phosphorus levels. Canonical Correspondence Analysis identified the perennial non-native grasses of high biomass as species most dependent on high soil nutrient levels. Such species may be resource-limited in undisturbed soils. Three classes of non-native plants have invaded this species-rich grassland: (1) generalist species (> 50 % frequency), mostly therophytes with non-specialized habitat or germination requirements; (2) resource-limited species comprising perennial species of high biomass that are dependent on nutrient increases and/or soil disturbances before they can invade the community and; (3) species of intermediate frequency (1–30 %), of low to high biomass potential, that appear to have non-specialized habitat requirements but are currently limited by seed dispersal, seedling establishment or the current site management. Native species richness and cover are most negatively affected by increases in non-native cover. Declines are largely evident once the non-native cover exceeds 40 %. Widespread, generalist non-native species are numerous in intact sites and will have to be considered a permanent part of the flora of remnant grasslands. Management must aim to minimize increases in cover of any non-native species or the disturbances that favour the establishment of competitive non-native grasses if the native grassland flora is to be conserved in small, fragmented remnants.     

Factors determining the centrifugal organization of remnant Festuca grassland communities in Alberta

Abstract. This paper describes the species composition of remnant grasslands in the aspen parkland region of Alberta and its relation to soil characteristics and small‐scale disturbance. Our findings are consistent with the centrifugal model of communities with Festuca hallii dominating undisturbed ‘core’ habitat and the composition of more ‘peripheral’ habitats varying in soil properties and in the magnitude of disturbance. Invasive non‐native species are not present in the core habitat and are present only in the disturbed sites, most abundantly in those with the highest soil nitrogen. The centrifugal model, as it applies to these remnant grasslands, differs from its previous application to wetlands and forests in that the core communities are not on the most fertile sites, but on the least disturbed. These findings have implications for the management of prairie remnants to exclude invasive exotic species.     

Inoculation with native soil improves seedling survival and reduces non-native reinvasion in a grassland restoration

Losses of grasslands have been largely attributed to widespread land-use changes, such as conversion to row-crop agriculture. The remaining tallgrass prairie faces further losses due to biological invasions by non-native plant species, often with resultant ecosystem degradation. Of critical concern for conservation, restoration of native grasslands has been met with little success following eradication of non-native plants. In addition to the direct and indirect effects of non-native invasive plants on beneficial soil microbes, management practices targeting invasive species may also negatively affect subsequent restoration efforts. To assess mechanisms limiting germination and survival of native species and to improve native species establishment, we established six replicate plots of each of the following four treatments: (1) inoculated with freshly collected prairie soil with native seeds; (2) inoculated with steam-pasteurized soil with native seeds; (3) noninoculated with native seeds; or (4) noninoculated/nonseeded control. Inoculation with whole soil did not improve seed germination; however, addition of whole soil significantly improved native species survival, compared to pasteurized soil or noninoculated treatments. Inoculation with whole soil significantly decreased reestablishment of non-native invasive Bothriochloa bladhii (Caucasian bluestem); at the end of the growing season, plots receiving whole soil consisted of approximately 30% B. bladhii cover, compared to approximately 80% in plots receiving no soil inoculum. Our results suggest invasion and eradication efforts negatively affect arbuscular mycorrhizal hyphal and spore abundances and soil aggregate stability, and inoculation with locally adapted soil microbial communities can improve metrics of restoration success, including plant species richness and diversity, while decreasing reinvasion by non-native species.     

Interactions between annuals and woody perennials in a Western Australian nature reserve

Abstract. We studied the interactions between woody perennial species and native and non-native annual species in a number of vegetation types within a nature reserve in the Western Australian wheatbelt. In particular, we examined the responses of annuals to perennial canopy removal, fire, soil disturbance and nutrient additions, and the effects of removal of annuals on perennial seedling regeneration. Experimental shrub removal significantly increased the abundance of annuals in a dense shrubland dominated by Allocasuarina campestris, but had no effect in a more open species-rich sandplain heath. Soil disturbance and nutrient addition in the heath area had no significant influence on annual abundance until three years after treatment. Fire had no clear effect on annual abundance in the heath within the reserve, but promoted a large increase in non-native species within an adjacent roadverge. A pattern of increased soil nutrient levels was accompanied by greatly increased non-native annual abundance beneath individual trees of Santalum spicatum. Exploratory laboratory bioassay experiments indicated that several woody perennials produced leachates that were capable of reducing the germination or growth of the introduced grass Avena fatua, indicating that allelopathy may be an important component of the interaction between the annual and perennial components. Within a woodland community, fire temporarily reduced the abundance of annual species and increased the establishment of perennial seedlings. Field experiments showed that annuals significantly reduced the survival of seedlings of the shrub Allocasuarina campestris. Our results indicate that intact native vegetation canopies effectively prevent invasion by non-native annuals, and that regeneration by native perennials is likely to be inhibited by the presence of an abundant annual cover.     

The influence of aqueous extracts of burnt or heated soil on the activity of vesicular-arbuscular mycorrhizal fungi propagules

Summary Aqueous extracts of burnt soil, unburnt soil and oven-heated unburnt soil were tested as to their effects on vesicular-arbuscular mycorrhizal (VAM) fungi (spore germination, mycelial propagule activity and root colonization). The extracts of burnt or heated soil inhibited VAM spore germination and extrarrhizal mycelium activity.