Fire-driven alien invasion in a fire-adapted ecosystem

Disturbance plays a key role in many alien plant invasions. However, often the main driver of invasion is not disturbance per se but alterations in the disturbance regime. In some fire-adapted shrublands, the community is highly resilient to infrequent, high-intensity fires, but changes in the fire regime that result in shorter fire intervals may make these communities more susceptible to alien plant invasions. This study examines several wildfire events that resulted in short fire intervals in California chaparral shrublands. In one study, we compared postfire recovery patterns in sites with different prefire stand ages (3 and 24 years), and in another study we compared sites that had burned once in four years with sites that had burned twice in this period. The population size of the dominant native shrub Adenostoma fasciculatum was drastically reduced following fire in the 3-year sites relative to the 24-year sites. The 3-year sites had much greater alien plant cover and significantly lower plant diversity than the 24-year sites. In a separate study, repeat fires four years apart on the same sites showed that annual species increased significantly after the second fire, and alien annuals far outnumbered native annuals. Aliens included both annual grasses and annual forbs and were negatively correlated with woody plant cover. Native woody species regenerated well after the first fire but declined after the second fire, and one obligate seeding shrub was extirpated from two sites by the repeat fires. It is concluded that some fire-adapted shrublands are vulnerable to changes in fire regime, and this can lead to a loss of native diversity and put the community on a trajectory towards type conversion from a woody to an herbaceous system. Such changes result in alterations in the proportion of natives to non-natives, changes in functional types from deeply rooted shrubs to shallow rooted grasses and forbs, increased fire frequency due to the increase in fine fuels, and changes in carbon storage.     

Management strategies for plant invasions: manipulating productivity, disturbance, and competition

The traditional approach to understanding invasions has focused on properties of the invasive species and of the communities that are invaded. A well‐established concept is that communities with higher species diversity should be more resistant to invaders. However, most recently published field data contradict this theory, finding instead that areas with high native plant diversity also have high exotic plant diversity. An alternative environment‐based approach to understanding patterns of invasions assumes that native and exotic species respond similarly to environmental conditions, and thus predicts that they should have similar patterns of abundance and diversity. Establishment and growth of native and exotic species are predicted to vary in response to the interaction of plant growth rates with the frequency and intensity of mortality‐causing disturbances. This theory distinguishes between the probability of establishment and the probability of dominance, predicting that establishment should be highest under unproductive and undisturbed conditions and also disturbed productive conditions. However, the probability of dominance by exotic species, and thus of potential negative impacts on diversity, is highest under productive conditions. The theory predicts that a change in disturbance regime can have opposite effects in environments with contrasting levels of productivity. Manipulation of productivity and disturbance provides opportunities for resource managers to influence the interactions among species, offering the potential to reduce or eliminate some types of invasive species.     

火与外来植物相互关系的研究进展

入侵种对本地的生态系统和生物多样性均有不良的影响,严重的会造成物种的灭绝和生态系统的崩溃,这已在全球范围引起广泛关注.在植物外来种与火生态因子的作用研究中发现,火与外来种的关系随物种生物学特性、火作用的时间、频度、强度不同而不同,火有时会有效地抑制外来种的生长和入侵,有时会促进一些外来种的生长和入侵.反之,一些外来种会对火的产生起到积极的作用,一些外来种又会抑制火的发生.火作为控制入侵种的一种方法,经科学地运用,可对某些入侵种起到有效的控制作用.     

Impacts of changed fire regimes on tropical riparian vegetation invaded by an exotic vine

Abstract Riparian habitats are highly important ecosystems for tropical biodiversity, and highly threatened ecosystems through changing disturbance regimes and weed invasion. An experimental study was conducted to assess the ecosystem impacts of fire regimes introduced for the removal of the exotic woody vine, Cryptostegia grandiflora, in tropical north‐eastern Australian woodlands. Experimental sites in subcatchments of the Burdekin River, northern Queensland, Australia, were subjected to combinations of early wet‐season and dry‐season fires, and single and repeated fires, with an unburnt control. Woody vegetation was sampled using permanent quadrats to record and monitor plants species, number and size‐class. Sampling was conducted pre‐fire in 1999 and post‐fire in 2002. All fire regimes were effective in reducing the number and biomass of C. grandiflora shrubs and vines. Few woodland or riparian species were found to be fire‐sensitive and community composition did not change markedly under any fire regime. The more intense dry‐season fires impacted the structure of non‐target vegetation, with large reductions in the number of sapling trees (<5 cm d.b.h.) and reductions in the largest tree size‐class and total tree basal area. Unexpectedly, medium‐sized canopy trees (10–30 cm d.b.h.) appear to have been significantly benefited by fires, with decreases in number of trees of this size‐class in the absence of fire. Although the presence of C. grandiflora as a vine in riparian forest canopies changed the nature and intensity of crown combustion patterns, this did not lead to the initiation of a self‐perpetuating weed–fire cycle, as invaders were unable to take advantage of gaps caused by fire. Low intensity, early wet‐season burning, or early dry‐season burning, is recommended for control of C. grandiflora in order to minimize the fire intensity and risk of the loss of large habitat trees in riparian habitats.     

Positive frequency dependence undermines the success of restoration using historical disturbance regimes

Anthropogenic alterations of historical disturbance regimes (e.g. suppressing floods and wildfires) is a primary mechanism by which exotic species can come to dominate native communities. Unfortunately, reinstating historical disturbance regimes to restore native communities has achieved mixed success. The presence of positive frequency dependence (PFD) is commonly invoked to explain why exotic plant invasions are so difficult to eradicate. However, models examining PFD have not considered the effect of reintroducing disturbances. Using a spatially explicit individual‐based model, we consider how magnitude and direction of frequency dependence of native and exotic species affects the success of reintroducing disturbances that favour fitness of natives over exotics. Our model illustrates why restoration is difficult; there is a narrow range of parameters that allows for native species to eliminate or coexist with exotics once they have established. Dominance by exotic invaders occurs with moderate initial frequencies of exotic individuals, aggregation of these individuals, or an exotic propagule production advantage. Reintroducing disturbances allows native dominance only when PFD of the exotic is weaker than that of the native species, disturbance intervals are short, and/or exotics are not initially frequent. Our framework provides guidelines for conditions in which the reintroduction of disturbances will effectively restore invaded habitats.     

Exotic herbivores and fire energy drive standing herbaceous biomass but do not alter compositional patterns in a semiarid savanna ecosystem

Questions

Fire regime alterations are pushing open ecosystems worldwide past tipping points where alternative steady states characterized by woody dominance prevail. This reduces the frequency and intensity of surface fires, further limiting their effectiveness for controlling cover of woody plants. In addition, grazing pressure (exotic or native grazers) can reinforce woody encroachment by potentially reducing fine-fuel loads. We investigated the effects of different fire energies on the herbaceous plant community, together with mammalian wildlife herbivory (exotic and native combined) exclusion, to inform best management practices.

Location

Texas semi-arid savanna, southern Great Plains, USA.

Methods

We conducted an experiment in which we manipulated fire intensity and herbivore access to herbaceous biomass in a split-plot design. We altered fire energy via fuel addition rather than applying fire under different environmental conditions to control for differences in standing biomass and composition attributable to differential plant physiological status and fire season.

Results

High-energy fire did not reduce herbaceous biomass or alter plant community composition, although it did increase among-plot variability in composition and forb biomass relative to low-energy fire and non-burned controls. Grazing pressure from native and non-native mammalian herbivores reduced above-ground herbaceous biomass regardless of fire treatments, but did not alter community composition.

Conclusions

Managers seeking to apply high-intensity prescribed fire to reduce woody encroachment will not negatively impact herbaceous plant productivity or alter community composition. However, they should be cognizant that repeated fires necessary for greatly reducing woody plants in heavily invaded areas might be difficult to accomplish due to fine-fuel reduction from wild herbivores. High fencing to restrict access by wildlife herbivores or culling might be necessary to build fuels sufficient to conduct high-intensity burns for woody-plant reduction.     

Positive interaction between invasive plants: The influence of Pyracantha angustifolia on the recruitment of native and exotic woody species

Abstract: Positive interactions between species are known to play an important role in the dynamics of plant communities, including the enhancement of invasions by exotics. We studied the influence of the invasive shrub Pyracantha angustifolia (Rosaceae) on the recruitment of native and exotic woody species in a secondary shrubland in central Argentina mountains. We recorded woody sapling recruitment and micro‐environmental conditions under the canopies of Pyracantha and the dominant native shrub Condalia montana (Rhamnaceae), and in the absence of shrub cover, considering these situations as three treatments. We found that native and exotic species richness were higher under Pyracantha than under the other treatments. Ligustrum lucidum (Oleaceae), an exotic bird‐dispersed shade‐tolerant tree, was the most abundant species recruiting in the area, and its density was four times higher under the canopy of Pyracantha. This positive interaction may be related to Pyracantha's denser shading, to the mechanical protection of its canopy against ungulates, and/or to the simultaneous fruit ripening of both woody invaders.     

Invasive belowground mutualists of woody plants

Most plants require mutualistic associations to survive, which can be an important limitation on their ability to become invasive. There are four strategies that permit plants to become invasive without being limited by a lack of mutualists. One is to not be dependent on mutualists. The other three strategies are to form novel mutualisms, form associations with cosmopolitan species, or co-invade with mutualists from their native range. Historically there has been a bias to study mutualisms from a plant perspective, with little consideration of soil biota as invasive species in their own right. Here we address this by reviewing the literature on belowground invasive mutualists of woody plants. We focus on woody invaders as ecosystem-transforming plants that frequently have a high dependence on belowground mutualists. We found that co-invasions are common, with many ectomycorrhizal plant species and N-fixing species co-invading with their mutualists. Other groups, such as arbuscular mycorrhizal plants, tend to associate with cosmopolitan fungal species or to form novel associations in their exotic range. Only limited evidence exists of direct negative effects of co-invading mutualists on native mutualist communities, and effects on native plants appear to be largely driven by altered environmental conditions rather than direct interactions. Mutualists that introduce novel ecosystem functions have effects greater than would be predicted based solely on their biomass. Focusing on the belowground aspects of plant invasions provides novel insights into the impacts, processes and management of invasions of both soil organisms and woody plant species.     

Native granivores reduce the establishment of native grasses but not invasive <Emphasis Type="Italic">Bromus tectorum</Emphasis>

Seed predation can structure plant communities by imposing strong population controls on some species but not others. In this context, studies from various ecosystems report that native granivores selectively forage for seeds from native species over seeds from exotic invaders, which could disproportionately favor the establishment of invaders and facilitate their dominance in communities. However, few studies have connected selective foraging for native seeds to differential patterns of establishment among native and invasive species. Thus, the extent to which preferential foraging for native seeds favors the establishment of invasive plants is unclear. Here, we used experimental seed additions and exclosure treatments at five field sites distributed across?≈?80,000 km² of the Great Basin Desert, USA to compare the effects of rodent foraging on the establishment of less-preferred cheatgrass (Bromus tectorum—an annual species native to Eurasia that is exotic and highly invasive across the Great Basin) and four species of more-preferred native grasses that commonly co-occur with cheatgrass. Rodent foraging reduced the establishment of each native species by at least 80% but had no effect on the establishment of cheatgrass, and this finding was consistent across study sites. Our results suggest that selective foraging for native species may favor the establishment of cheatgrass over native grasses, potentially exacerbating one of the most extensive plant invasions in North America.     

Ecological filtering of exotic plants in an Australian sub‐alpine environment

We investigated some of the factors influencing exotic invasion of native sub‐alpine plant communities at a site in southeast Australia. Structure, floristic composition and invasibility of the plant communities and attributes of the invasive species were studied. To determine the plant characteristics correlated with invasiveness, we distinguished between roadside invaders, native community invaders and non‐invasive exotic species, and compared these groups across a range of traits including functional group, taxonomic affinity, life history, mating system and morphology. Poa grasslands and Eucalyptus‐Poa woodlands contained the largest number of exotic species, although all communities studied appeared resilient to invasion by most species. Most community invaders were broad‐leaved herbs while roadside invaders contained both herbs and a range of grass species. Over the entire study area the richness and cover of native and exotic herbaceous species were positively related, but exotic herbs were more negatively related to cover of specific functional groups (e.g. trees) than native herbs. Compared with the overall pool of exotic species, those capable of invading native plant communities were disproportionately polycarpic, Asteracean and cross‐pollinating. Our data support the hypothesis that strong ecological filtering of exotic species generates an exotic assemblage containing few dominant species and which functionally converges on the native assemblage. These findings contrast with those observed in the majority of invaded natural systems. We conclude that the invasion of closed sub‐alpine communities must be viewed in terms of the unique attributes of the invading species, the structure and composition of the invaded communities and the strong extrinsic physical and climatic factors typical of the sub‐alpine environment.     

Mechanisms underlying the impacts of exotic plant invasions

Although the impacts of exotic plant invasions on community structure and ecosystem processes are well appreciated, the pathways or mechanisms that underlie these impacts are poorly understood. Better exploration of these processes is essential to understanding why exotic plants impact only certain systems, and why only some invaders have large impacts. Here, we review over 150 studies to evaluate the mechanisms underlying the impacts of exotic plant invasions on plant and animal community structure, nutrient cycling, hydrology and fire regimes. We find that, while numerous studies have examined the impacts of invasions on plant diversity and composition, less than 5% test whether these effects arise through competition, allelopathy, alteration of ecosystem variables or other processes. Nonetheless, competition was often hypothesized, and nearly all studies competing native and alien plants against each other found strong competitive effects of exotic species. In contrast to studies of the impacts on plant community structure and higher trophic levels, research examining impacts on nitrogen cycling, hydrology and fire regimes is generally highly mechanistic, often motivated by specific invader traits. We encourage future studies that link impacts on community structure to ecosystem processes, and relate the controls over invasibility to the controls over impact.     

Plant invasions differentially affected by diversity and dominant species in native‐ and exotic‐dominated grasslands

Plant invasions are an increasingly serious global concern, especially as the climate changes. Here, we explored how plant invasions differed between native‐ and novel exotic‐dominated grasslands with experimental addition of summer precipitation in Texas in 2009. Exotic species greened up earlier than natives by an average of 18 days. This was associated with a lower invasion rate early in the growing season compared to native communities. However, invasion rate did not differ significantly between native and exotic communities across all sampling times. The predictors of invasion rate differed between native and exotic communities, with invasion being negatively influenced by species richness in natives and by dominant species in exotics. Interestingly, plant invasions matched the bimodal pattern of precipitation in Temple, Texas, and did not respond to the pulse of precipitation during the summer. Our results suggest that we will need to take different approaches in understanding of invasion between native and exotic grasslands. Moreover, with anticipated increasing variability in precipitation under global climate change, plant invasions may be constrained in their response if the precipitation pulses fall outside the normal growing period of invaders.     

Understory response to varying fire frequencies after 20 years of prescribed burning in an upland oak forest

Ecosystems in the eastern United States that were shaped by fire over thousands of years of anthropogenic burning recently have been subjected to fire suppression resulting in significant changes in vegetation composition and structure and encroachment by invasive species. Renewed interest in use of fire to manage such ecosystems will require knowledge of effects of fire regime on vegetation. We studied the effects of one aspect of the fire regime, fire frequency, on biomass, cover and diversity of understory vegetation in upland oak forests prescribe-burned for 20 years at different frequencies ranging from zero to five fires per decade. Overstory canopy closure ranged from 88 to 96% and was not affected by fire frequency indicating high tolerance of large trees for even the most frequent burning. Understory species richness and cover was dominated by woody reproduction followed in descending order by forbs, C3 graminoids, C4 grasses, and legumes. Woody plant understory cover did not change with fire frequency and increased 30% from one to three years after a burn. Both forbs and C3 graminoids showed a linear increase in species richness and cover as fire frequency increased. In contrast, C4 grasses and legumes did not show a response to fire frequency. The reduction of litter by fire may have encouraged regeneration of herbaceous plants and helped explain the positive response of forbs and C3 graminoids to increasing fire frequency. Our results showed that herbaceous biomass, cover, and diversity can be managed with long-term prescribed fire under the closed canopy of upland oak forests.     

Post‐fire vegetation dynamics in nutrient‐enriched and non‐enriched sclerophyll woodland

Abstract Exotic plant invasions are a significant problem in urban bushland in Sydney, Australia. In low‐nutrient Hawkesbury Sandstone communities, invasive plants are often associated with urban run‐off and subsequent increases in soil nutrients, particularly phosphorus. Fire is an important aspect of community dynamics in Sydney vegetation, and is sometimes used in bush regeneration projects as a tool for weed control. This study addressed the question: ‘Are there differences in post‐fire resprouting and germination of native and exotic species in nutrient‐enriched communities, compared with communities not disturbed by nutrient enrichment?’ We found that in non‐enriched areas, few exotic species emerged, and those that did were unable to achieve the rapid growth that was seen in exotic plants in the nutrient‐enriched areas. Therefore, fire did not promote the invasion of exotic plants into areas that were not nutrient‐enriched. In nutrient‐enriched areas after fire, the diversity of native species was lower than in the non‐enriched areas. Some native species were able to survive and compete with the exotic species in terms of abundance, per cent cover and plant height. However, these successful species were a different suite of natives to those commonly found in the non‐enriched areas. We suggest that although fire can be a useful tool for short‐term removal of exotic plant biomass from nutrient‐enriched areas, it does not promote establishment of native species that were not already present.     

Fire indirectly benefits fitness in two invasive species

Ecosystems perturbed from their natural disturbance regimes are more vulnerable to establishment and dominance of exotic plant species. Restoration efforts that reintroduce fire have achieved mixed success in reducing the abundance of exotic plants. The responses of many native species to fire are well known; fire-adapted species respond directly (heat and smoke cue germination) and indirectly (post-fire environment benefits seedling survivorship and growth) to fire. However, the direct and indirect effects of fire are unknown for most exotic plant species. We tested the direct and indirect effects of fire on two exotic invaders of Asian origin, Ailanthus altissima and Lonicera maackii, in North American woodlands. To quantify the direct effects of fire, we compared germination rates of seeds exposed to varying levels of heat and smoke in a laboratory and placed at different soil depths during a prescribed fire in the field. We examined the indirect effects of fire by comparing seedling recruitment in burned and unburned woodland plots. Results indicate that neither A. altissima nor L. maackii have germination cues associated with fire. However, both species have greater seedling recruitment in burned as compared to unburned areas in the field. Although seeds of these invasive species are not specifically adapted to fire, they still benefit from post-fire environments and pose a challenge to restoration of fire-maintained ecosystems. Future studies using our approach will allow land managers to better predict how communities will respond to restoration efforts and to understand variability observed in past restoration projects.     

The role of blackbirds (

The naturalised European blackbird (Turdus merula) is the most widely distributed avian seed disperser in New Zealand. Together with the native silvereye (Zosterops lateralis) they are the major seed dispersers over large areas of New Zealand. I review the international literature on aspects of the ecology and behaviour of blackbirds relevant to their potential for dispersing weeds in New Zealand. Blackbirds eat a wide range of native and exotic fruit including many naturalised species. Their habitat preferences and behaviour result in germinable seeds being deposited in a range of sites, particularly in shrubby habitats, where seedling establishment is likely. Most seeds will be deposited within 50 m but some may be carried a kilometre or more to develop new invasive loci. Blackbirds therefore probably make a major contribution to the development of novel plant communities of naturalised woody weeds. These communities provide fruit more suited to non- endemic native birds and naturalised birds, than to endemic birds. The relative contribution of blackbirds and silvereyes to seed dispersal of native and exotic species requires investigation. The outcome may suggest potential for managing blackbirds as a vector of weed invasions.     

Influence of woody invader control methods and seed availability on native and invasive species establishment in a Hawaiian forest

When invasive woody plants become dominant, they present an extreme challenge for restoration of native plant communities. Invasive Morella faya (fire tree) forms extensive, nearly monospecific stands in wet and mesic forests on the Island of Hawai’i. We used logging, girdling, and selective girdling over time (incremental girdling) to kill stands of M. faya at different rates, with the objective of identifying a method that best promotes native forest re-establishment. We hypothesized that rapid canopy opening by logging would lead to establishment of fast-growing, non-native invaders, but that slower death of M. faya by girdling or incremental girdling would increase the establishment by native plants adapted to partial shade conditions. After applying the M. faya treatments, seed banks, seed rain, and plant recruitment were monitored over 3 years. Different plant communities developed in response to the treatments. Increased light and nitrogen availability in the logged treatment were associated with invasion by non-native species. Native species, including the dominant native forest tree, (Metrosideros polymorpha) and tree fern (Cibotium glaucum), established most frequently in the girdle and incremental girdle treatments, but short-lived non-native species were more abundant than native species. A diverse native forest is unlikely to develop following any of the treatments due to seed limitation for many native species, but girdling and incremental girdling promoted natural establishment of major components of native Hawaiian forest. Girdling may be an effective general strategy for reestablishing native vegetation in areas dominated by woody plant invaders.     

Fire as a Restoration Tool: A Decision Framework for Predicting the Control or Enhancement of Plants Using Fire

Wildfires change plant communities by reducing dominance of some species while enhancing the abundance of others. Detailed habitat‐specific models have been developed to predict plant responses to fire, but these models generally ignore the breadth of fire regime characteristics that can influence plant survival such as the degree and duration of exposure to lethal temperatures. We provide a decision framework that integrates fire regime components, plant growth form, and survival attributes to predict how plants will respond to fires and how fires can be prescribed to enhance the likelihood of obtaining desired plant responses. Fires are driven by biotic and abiotic factors that dictate their temporal (seasonality and frequency), spatial (size and patchiness), and magnitude (intensity, severity, and type) components. Plant resistance and resilience to fire can be categorized by a combination of life form, size, and ability to disperse or protect seeds. We use a combination of life form and vital plant attributes along with an understanding of fire regime components to suggest a straightforward way to approach the use of fire to either reduce or enhance particular species. A framework for aiding decisions is organized by life form and plant size. Questions regarding perennating bud and seed characteristics direct restoration practitioners to fire regimes that may achieve their management objectives of either increasing or decreasing plants with specific life form characteristics.     

High seedling relative growth rate and specific leaf area are traits of invasive species: phylogenetically independent contrasts of woody angiosperms

Understanding causal factors of exotic species invasions is important not only for prevention and prioritizing control efforts, but also for providing valuable insights into the underlying biology of contrasting life-history strategies. In seedling growth analyses, invasive woody species were compared with less-invasive woody species commonly cultivated in California using phylogenetically corrected procedures (12 phylogenetically independent contrasts). Invasive species were hypothesized to have higher seedling relative growth rates (RGRs) and specific leaf areas (SLAs) than did related less-invasive species. In phylogenetically independent contrasts conducted among taxa within families, high seedling RGRs and SLAs have significant positive associations with woody plant invasiveness. For contrasts containing species invasive in mediterranean regions, invasive species had significantly larger root biomass allocation than did less-invasive species. Optimization of fast seedling growth (high RGR) associated with opportunistic resource acquisition (high SLA) and increased root allocation to survive summer drought may be critical for the success of plant invaders in regions with mediterranean climates.