Community changes following shrub invasion of grassland

Summary We studied the development of the shrub Baccharis pilularis ssp consanguinea and its effects on herbs of the annual grassland in Northern California. A series of shands of Baccharis was sampled of ages ranging from 1 yr to>9 yr, representing most of the life cycle of the shrub. In each stand we examined shrub biomass, structure and litterfall. We also determined cover and biomass of all herbaceous species and estimated seed production, seed rain and storage of seed in the soil. Abundances of all herbaceous species declined greatly after Baccharis formed a closed canopy at 2–3 yr, and little seed of herbaceous species was either dispersed into shrub stands or stored in the soil. Exclosures suggested that herbivory by small mammals in the closed shrub stands may be important in reducing the abundance of herbaceous species following shrub invasion of grassland.     

Unlocking the potential of Google Earth as a tool in invasion science

Distribution data are central to many invasion science applications. The shortage of good information on the distribution of alien species and their spatial dynamics is largely attributable to the cost, effort and expertise required to monitor these species over large areas. Virtual globes, particularly Google Earth, are free and user-friendly software which provide high-resolution aerial imagery for the entire globe. We suggest this has enormous potential for invasion science. We provide suggestions and tools for gathering data on the distribution and abundance of invasive alien trees using visual interpretation of Google Earth imagery, and propose how these data may be used for a number of purposes, including calculating useful metrics of invasions, prioritising species or areas for management and predicting potential distributions of species. We also suggest various practical uses of Google Earth, such as providing a tool for early detection of emerging invasions, monitoring invasions over time, and to help researchers and managers identify suitable field study sites. Virtual globes such as Google Earth are not without limitations and we provide guidance on how some of these can be overcome, or when imagery from Google Earth may not be fit for invasion science purposes. Because of Google Earth’s huge popularity and ease of use, we also highlight possibilities for awareness-raising and information sharing that it provides. Finally, we provide the foundations and guidelines for a virtual global network of sentinel sites for early detection, monitoring and data gathering of invasive alien trees, which we propose should be developed as part of a “citizen science” effort. There has been limited use of virtual globes by invasion scientists and managers; it is our hope that this paper will stimulate their greater use, both within the field of invasion science and within ecology generally.     

Long-term patterns of shrub expansion in a C4-dominated grassland: fire frequency and the dynamics of shrub cover and abundance

Worldwide, grassland ecosystems have experienced a major shift in growth-form dominance as woody plant species have expanded and replaced native grasses. In the C(4)-dominated grasslands of central North America, a reduction in fire frequency is the most cited cause of this shift in growth forms as fire both enhances grass productivity and constrains the establishment and expansion of native woody vegetation. Using an 18-yr plant species composition data set, we quantified patterns of change in shrub cover, frequency, and species richness associated with three distinct fire regimes. During the study period (1983-2000), shrub cover increased most dramatically in sites in which the frequency of fire was once every 4 yr (intermediate frequency; 28.6%) followed by sites in which fire occurred only once during the 18-yr period (low frequency; 23.7%). Annual fire effectively prevented the recruitment of new woody species, but even with this high fire frequency, shrub cover increased slightly (3.7%). Comparatively, shrub species richness increased by three and six, respectively, in the intermediate- and low-frequency fire sites. These data indicate that within this grassland, periods without fire are necessary for recruitment of both new individuals and additional shrub species; however, once established, shrub cover will increase regardless of fire frequency and even annual fire will not reduce shrub abundance.     

The effect of fire and ants on the seed-bank of a shrub in a semi-arid grassland

Abstract Invasion of grasslands by woody plants following the introduction of domestic stock is a worldwide phenomenon. Burning is frequently recommended as a remedial measure but for a pastoral enterprise it is costly and the frequency of the fires required is of critical economic importance. The size and longevity of the soil seed-bank is an essential part of the response of shrub populations to prescribed fire regimes. In this study the seed-bank of the semi-arid zone shrub Dodonaea attenuata in Eragrostis eriopoda tussock grassland was examined in relation to harvesting by ants and the burning history of the sites. On unburnt sites, more than 3500 seeds per m2 entered the seed-bank in the summer 1984–85 but sites burnt 5 years previously produced less than one-third of that number. Burnt shrubs did not flower for 5 years and no seeds survived in the soil through to 1985. Burning immediately prior to seed ripening destroyed the seed crop but burning after seed-fall stimulated a greater germination in the following spring than on unburnt treatments. Ants rapidly harvested most of the seeds produced and after 20 months the combined effects of ant harvesting and germination had reduced the seed-bank at unburnt sites to 8–21 seeds per m2 and at sites burnt 7 years previously to less than two seeds per m2. Seeds were initially harvested in summer by Pheidole spp. of ants for their elaiosome and then discarded in middens outside the entrance to the ant nest. During the subsequent cool season the seeds were taken back into the nest and stored at depths ranging from 2 to 30cm. It was concluded that ants provided short-range dispersal (< 10 m) and promoted the contagious distribution of D. attenuata, which is advantageous for a fire-susceptible, arid-zone shrub invading a grassland liable to be burnt: seedlings derived from seeds in ant storage chambers near the soil surface and in sparse grass situations caused by competition from shrubs may obtain some survival advantage. Prescribed fire has potential as a management tool for controlling population density of D. attenuata because, depending upon season and frequency, it reduces seed rain by killing shrubs, suppresses flowering activity and destroys seed crops on the plant. Under the influence of a regime of regular burning, such as prevailed prior to the European pastoral industry, the limited soil seed-bank would have been a major constraint on D. attenuata populations.     

Exotic shrub invasion in an undisturbed wetland has little community-level effect over a 15-year period

In this long-term study, we examined the invasion by the exotic shrub glossy buckthorn (Rhamnus frangula L.) and the response of co-occurring plants in a large, undisturbed wetland. We first sampled the vegetation in 1991 and repeated the sample 15 years later using the same, permanently located sample units (n = 165). Despite dramatic increases in the abundance of buckthorn, the invasion elicited little apparent response by the resident plant community. Species richness and cover in the herbaceous plant stratum had no apparent relationship with change in buckthorn cover. The number of shrub species other than buckthorn showed no relationship with change in buckthorn cover, but the cover of other shrubs decreased as buckthorn cover increased. Species composition changed independently of changes in buckthorn cover. These results show that dramatic increases in the abundance of an invasive species do not necessarily cause large changes in the native plant community and suggest disturbance history influences community response to invasion.     

Phylogenetic relatedness as a tool in restoration ecology: a meta-analysis

Biotic interactions assembling plant communities can be positive (facilitation) or negative (competition) and operate simultaneously. Facilitative interactions and posterior competition are among the mechanisms triggering succession, thus representing a good scenario for ecological restoration. As distantly related species tend to have different phenotypes, and therefore different ecological requirements, they can coexist, maximizing facilitation and minimizing competition. We suggest including phylogenetic relatedness together with phenotypic information as a predictor for the net effects of the balance between facilitation and competition in nurse-based restoration experiments. We quantify, by means of a Bayesian meta-analysis of nurse-based restoration experiments performed worldwide, the importance of phylogenetic relatedness and life-form disparity in the survival, growth and density of facilitated plants. We find that the more similar the life forms of neighbouring plants are the greater the positive effect of phylogenetic distance is on survival and density. This result suggests that other characteristics beyond life form are also contained in the phylogeny, and the larger the phylogenetic distance, the less is the niche overlap, and therefore the less is the competition. As a general rule, we can maximize the success of the nurse-based practices by increasing life-form disparity and phylogenetic distances between the neighbour and the facilitated plant.     

Allelopathy of a native grassland community as a potential mechanism of resistance against invasion by introduced plants

Successful plant invasions depend, at least partly, on interactions between introduced plants and native plant communities. While allelopathic effects of introduced invaders on native resident species have received much attention, the reverse, i.e. allelopathic effects of native residents on introduced plants, have been largely neglected. Therefore, we tested whether allelopathy of native plant communities decreases their invasibility to introduced plant species. In addition, we tested among the introduced species whether the invasive ones are more tolerant to allelopathy of native plant communities than the non-invasive ones. To test these hypotheses, we grew nine pairs of related (congeneric or confamilial) invasive and non-invasive introduced plant species (i.e. 18 species) in the presence or absence of a native grassland community, which consisted of three common forbs and three common grasses, with or without activated carbon in the soil. Activated carbon reduced the survival percentage and growth of introduced plants in the absence of the native plant community. However, its net effect on the introduced plants was neutral or even slightly positive in the presence of the native community. This might suggest that the native plant community imposed allelopathic effects on the introduced plants, and that these effects were neutralized or reduced by activated carbon. The invasive and non-invasive introduced plants, however, did not differ in their tolerance to such allelopathic effects of the native plant community. Thus, although allelopathy of native plant communities might increase their resistance against introduced plants, there was no evidence that tolerance to allelopathy of native plant communities contributes to the degree of invasiveness of introduced plants.     

Exotic grass invasion alters potential rates of N fixation in Hawaiian woodlands

Exotic grass invasion promotes fire which drives the conversion of native woodlands to exotic grasslands in the seasonally dry submontane forests of the island of Hawai'i. We compared potential rates of N fixation in an unburned forest site and a converted grassland site using the acetylene reduction assay. In addition to measuring rates of N fixation on separate and mixed substrates in each site, we tested the effect of abiotic factors on rates of N fixation of specific substrates. We hypothesized that rates of N fixation would be higher in the converted grassland site. N fixation estimates were 4.9 kg N ha⁻¹ year⁻¹ for the unburned forest, and 0.10 kg N ha⁻¹ year⁻¹ for the grassland site, so our hypothesis was rejected. The N fixation in the unburned forest occurs mostly on the leaf litter of native woody species. These substrates are absent from the grassland site, except for wood debris which was not consumed during the fires. No nitrogenase activity was detected in the rhizosphere and litter of grasses, the rhizospheres of shrubs or in soil. Although wood debris is not a significant contributor to the N fixed in the unburned forest, it contributes the majority of N fixed in the grassland. The response of nitrogenase activity to varying conditions of moisture and temperature suggests that microclimatic differences between sites do not control differences in N fixation activity; rather, these differences are due to the abundance of N-fixing substrates. The substantial decrease in N fixation activity after the conversion from woodland to grassland implies that ecosystem-level rates of N accretion are decreased by fire in these sites so much that the N lost during volatilization due to fire is not replenished over the long term by N fixation. Received: 10 January 1997 / Accepted: 7 August 1997     

Shrub succession and invasibility in a New Zealand montane grassland

Abstract Two successive shrub invasions of a short tussock grassland induced by grazing and burning were examined in montane South Island, New Zealand. The first invasion was by a native shrub, matagouri (Discaria toumatou Raoul). The second invasion was by an exotic shrub, Scotch broom (Cytisus scoparius (L.) Link), which invaded the matagouri shrubland that had developed over the grassland. The invasions were investigated using analysis of spatial patterns of both shrubs and tussocks, and age, growth rates and size structure of the shrubs. Competition between the two shrub species was examined using spatial patterns and comparing allometric relationships. After initial invasion by matagouri of the grasslands, stand density increased by consolidation about its initial colonization points. Current matagouri distribution is often negatively associated with tussocks. Scotch broom occurs most frequently in a dense sward of introduced grasses and occasionally in tussocks in interstices among matagouri shrubs. Despite the palatability of Scotch broom to sheep that graze the site, there was no evidence that the spiny matagouri facilitates invasion by protecting Scotch broom seedlings; rather there was negative association between the shrub species. The two species probably compete for above-ground space. However, diameter and height growth rates of Scotch broom far exceed those of matagouri so Scotch broom is likely to increase in biomass rapidly at the site. The autogenic organization and disturbance history of the resident plant communities have rendered each vulnerable to successive invasions.     

The potential role of public gardens as sentinels of plant invasion

Public gardens can help prevent detrimental effects of plant invasions by collecting and sharing data on taxa spreading from cultivation early in the invasion process, thereby acting as sentinels of plant invasion. Existing initiatives have called for public gardens to adopt measures preventing plant invasion, but it is unclear what actions individual gardens are implementing, as there is no formal mechanism for communicating their progress. This study used internal lists of escaping taxa from seven public gardens in the Midwestern United States and Canada to demonstrate how public gardens can collectively contribute data that is critical to assessing potential invasiveness. It also reveals methodological differences in how gardens develop their lists of escaping plants, leading to recommendations for standardization. Data pooled across gardens yielded 769 species spreading from cultivation at one or more gardens. Eight woody species were listed by all gardens despite not consistently being recognized as invasive by states and provinces containing the gardens; some species recorded by multiple gardens did not appear on any invasive lists. While it may be premature to call taxa escaping from cultivation at a few public gardens “invasive” or even “potentially invasive”, these plants should be monitored and evaluated with this information shared to facilitate stronger conclusions about risk. Thus, public gardens have a unique expertise in assisting invasive plant efforts as sentinels, particularly if challenges related to methodological inconsistencies and data sharing are suitably addressed, which is herein recommended through the adoption of a set of standardized guidelines.

     

Correction to: The potential role of public gardens as sentinels of plant invasion

Biodiversity and Conservation -     

Assessment of interaction between sheep and poorly palatable grass: a key tool for grassland management and restoration

Invasion of poorly palatable grasses due to abandonment or improper grazing management decreases pastures feeding value and biodiversity. With the aim to control their spread, we assessed the relationship between sheep foraging behavior and changes in aboveground phytomass, leaf traits, and chemical features of the tall grass Brachypodium rupestre and evaluated the effects of a B. rupestre-based diet on epithelium keratinization of rumen. Our results demonstrated that sheep became less selective throughout the experimental trials and B. rupestre decreased its aboveground phytomass. Some leaf traits showed significant changes (LDMC, LA, and ADL were higher in ungrazed areas; leaf nitrogen content was higher in the grazed ones). In addition, we detected an increase of the degree of epithelium keratinization of sheep. Thus high grazing pressure can be used to control the spread of B. rupestre, but negative effects on animal welfare due to the increase of rumen keratinization might be expected.     

Gene-flow in the clouds: landscape genetics of a viviparous,montane grassland toad in the tropics

Anthropogenic habitat alteration often increases fragmentation and isolation, which decreases population sizes and increases extinction risk for species. Extrinsic threats may be buffered or enhanced by intrinsic factors. Within amphibians, the influence of different environmental and intrinsic factors on the population structure is not yet fully understood. Four factors were found to be important for population connectivity: life history traits, recent (anthropogenic) land use history, habitat, and topography, but the direction of their influence differed between studies. Here, we examine the genetic population structure and interpopulation connectivity within the complete distribution of Nimba toads (Nimbaphrynoides occidentalis), a toad from montane tropical West Africa. The Nimba toad is the only known viviparous, matrotrophic (foetuses are nourished during the gestation by their mothers) anuran on Earth. It occurs in three regions, the smallest is situated in disturbed, the largest population in partly disturbed habitat and the third was not yet impacted. We found small, but significant population differentiation, no indication of a recent bottleneck in the smallest population, but an indication of a reduction in population sizes in the more distant past in all three populations and no sex-biased dispersal. Correlations with landscape classifications indicate that high elevations, due to their high humidity levels, are the most important landscape characteristic facilitating dispersal. This underscores desiccation risk as an important landscape characteristic for amphibian population connectivity. We found indication that life-history traits (viviparity), land use history (mining-related activity) and topography (elevation) have an influence on Nimba toad population differentiation and gene-flow.     

Urokinase directly activates matrix metalloproteinases-9: a potential role in glioblastoma invasion

Previous reports showed that urokinase plasminogen activator (uPA) converts plasminogen to plasmin which then activates matrix metalloproteinases (MMPs). Here, we report that uPA directly cleaved pro-MMP-9 in a time-dependent manner at both C- and N-terminus and generated two gelatinolytic bands. uPA-activated-MMP-9 efficiently degraded fibronectin and blocked by uPA inhibitor B428 and recombinant tissue inhibitor of metalloproteinase-1 (TIMP-1). B428 inhibited basal and PMA-induced active MMP-9 in glioblastomas (GBM) U1242 cell media as well as cell invasion in vitro. A combination of MMP-9 and uPA antibodies more significantly inhibited U1242 cell invasion than uPA or MMP-9 antibody alone. Both uPA and MMP-9 were highly expressed in U1242 cell and GBM patient specimens. Furthermore, two active MMP-9 fragments with identical molecular weights to the uPA-activated MMP-9 products were detected in GBM patient specimens. These results suggest that uPA-mediated direct activation of MMP-9 may promote GBM cell invasion.     

Observations of responses to re‐introducing fire in a Basalt Plains grassland after the removal of grazing: Implications for restoration

Natural grasslands in southern Australia commonly exist in altered states. One widespread altered state is grassland pasture dominated by cool‐season (C3) native grasses maintained by ongoing grazing. This study explores the consequences of removing grazing and introducing fire as a conservation management tool for such a site. We examined the abundance of two native and three exotic species, across a mosaic of fire regimes that occurred over a three‐year period: unburnt, summer wild‐fire (>2 years previous), autumn management fire (<1 year previously) and burnt in both fires. Given that one aim of conservation management is to increase native species at the expense of exotics, the impacts of the fires were largely positive. Native grasses were at higher cover levels in the fire‐managed vegetation than in the unburnt vegetation. Of the three exotic species, one was consistently at lower density in the burnt plots compared to the unburnt plots, while the others were lower only in those plots burnt in summer. The results show that the response of a species varies significantly between different fire events, and that the effects of one fire can persist through subsequent fires. Importantly, some of the effects were large, with changes in the density of plants of over 100‐fold. Fire is potentially a cost‐effective tool to assist the ecological restoration of retired grassland pastures at large scales.     

Amphibian species and functional diversity as indicators of restoration success in tropical montane forest

Tropical forest restoration is increasingly seen as an activity that may counteract or reduce biodiversity loss. However, few studies monitor fauna or consider measures of functional diversity to assess restoration success. We assessed the effect of a tropical montane forest restoration program on species and functional diversity, using amphibians as the target group. We compared amphibian assemblages in three types of land use: restoration areas, tropical montane cloud forest (TMCF; reference ecosystem) and cattle pastures (degraded ecosystem) in southern Mexico. We also described microclimate, microhabitat heterogeneity, woody vegetation structure and diversity for each type of land use, and their relationship to amphibian species and functional diversity. Compared to TMCF, restoration areas had similar environmental conditions. However, amphibian species richness was similar in the three types of land use and abundance was lower in the restoration areas. In TMCF, the amphibian assemblage was dominated by forest-specialist species, the pastures by generalist species, and the restoration areas by a combination of both species types. Interestingly, functional richness, functional evenness and functional divergence did not vary with land use, though the number of functional groups in restoration areas and TMCF was slightly higher. Overall, the results suggest that after seven years, active restoration provided habitat heterogeneity and recovered woody vegetation capable of maintaining amphibian species and functional groups similar to those inhabiting TMCF. Forest fragments adjacent to restoration areas seem to facilitate fauna recolonization and this emphasizes the importance of the conservation of the reference ecosystems to achieving restoration success.     

Phytosociology as a tool for forest restoration: a study case in the extreme South of Atlantic Forest Biome

The biodiversity loss, due to anthropic activities, has generated the necessity of studies that enable the ecosystem conservation. Nevertheless, the majority of studies restrict themselves to the floristic composition and phytosociological structure from determined pre-established diameter, neglecting attributes of the species, such as its (their) niche (s) of occurrence in the forest, thus, making it difficult to identify potential species for restoration. In this way, the present study aimed to evaluate the structure of a forest fragment in the extreme South of Atlantic Forest Biome, through identification of species in the different layers. There were used 18 parcels of 10 m × 20 m, distributed systematically, where the vegetation was evaluated in four size classes. The characterization of the structure of the arboreal-shrub of the forest was performed by the data comparison of the floristic composition and phytosociological structure. In the fragment of forest studied, there were sampled a total of 2.176 individuals. The floristic composition of the area includes 74 species, belonging to 58 genera and 28 botanical families. We observed differences in relation of richness and diversity of species among the analyzed layers, the intermediate layer presented the lowest richness of species, being the pioneers and initial secondaries the least representatives. Trichilia claussenii, Sorocea bonplandii, Actinostemon concolor, Apuleia leiocarpa and Nectandra megapotamica were the most abundant species and well distributed in the different size classes, and they can be used in the enrichment of areas in process of restoration in the scope of the Deciduous Seasonal Forest in the extreme South of Atlantic Forest Biome. Ocotea puberula, Allophylus edulis, Cedrela fissillis and Machaerium paraguariense formed a group of species with expressive representativeness in the arboreal layer, thus they can be used in the moment of planting as species to the fast covering of the area.