Impacts of extreme climatic events on competition during grassland invasions

Phytometers of five C3 and five C4 species were transplanted into three different grasslands to study the effects of extreme climatic events on community invasibility and competition. Single extreme heating (eight hours at 52.5 °C) and rainfall (the equivalent of 100 mm) events in factorial combinations were superimposed on the grassland communities. A novel technique involving portable computer‐controlled chambers was used to create the heating events. In order to generate predictions of response to the extreme climatic events, the 10 phytometer species were categorized on the basis of 12 key plant functional traits. Using principal component analysis, two functional types (FTs) were identified as most likely to be advantaged (FT1, fast‐growing C4 annuals) and disadvantaged (FT2, slower‐growing C3 perennials) by an extreme climatic event. Competition between the resident vegetation and FT1 plus other C4 phytometers was consistently more intense within the exclusively C3 community compared to the dry C3/C4 community or moist C3/C4 community. The single extreme heating event had the greatest impact on competition, lowering the intensity of competition between the phytometers and resident vegetation. Our results indicate that competition is highly important in limiting the invasion of C3 grasslands by C4 species. The FT1 and FT2 responses confirmed predictions based on plant functional traits, whether growing as phytometers or as part of the resident vegetation. Future increases in climatic variability and the incidence of extreme climatic events are expected to suppress C3 competitive dominance and promote invasion of C4 species, in particular, the FT1 species.     

Native communities determine the identity of exotic invaders even at scales at which communities are unsaturated

Aim To determine why some communities are more invasible than others and how this depends on spatial scale. Our previous work in serpentine ecosystems showed that native and exotic diversity are negatively correlated at small scales, but became positively correlated at larger scales. We hypothesized that this pattern was the result of classic niche partitioning at small scales where the environment is homogeneous, and a shift to the dominance of coexistence mechanisms that depend on spatial heterogeneity in the environment at large scales. Location Serpentine ecosystem, Northern California. Methods We test the above hypotheses using the phylogenetic relatedness of natives and exotics. We hypothesized that (1) at small scales, native and exotic species should be more distantly related than expected from a random assemblage model because with biotic resistance, successful invaders should have niches that are different from those of the natives present and (2) at large scales, native and exotic species should not be more distantly related than expected. Result We find strong support for the first hypothesis providing further evidence of biotic resistance at small scales. However, at large scales, native and exotic species were also more distantly related than expected. Importantly, however, natives and exotics were more distantly related at small scales than they were at large scales, suggesting that in the transition from small to large scales, biotic resistance is relaxed but still present. Communities at large scales were not saturated in the sense that more species could enter the community, increasing species richness. However, species did not invade indiscriminately. Exotic species closely related to species already established the community were excluded. Main conclusions Native communities determine the identity of exotic invaders even at large spatial scales where communities are unsaturated. These results hold promise for predicting which species will invade a community given the species present.     

Effects of land use and environment on alien and native macrophytes: lessons from a large‐scale survey of Australian rivers

Aim We sampled riverine macrophyte communities and environmental conditions to compare drivers of alien and native abundance and to provide a general set of environmental correlates of invasion by aquatic macrophytes. Location Streams adjacent to three land‐use types (intensive, agricultural and natural) across a large latitudinal gradient (approximately 27° S–43° S) in Australia. Sites were located near Brisbane (Queensland), Sydney (New South Wales), Canberra (Australian Capital Territory), Melbourne (Victoria) and Hobart (Tasmania). Methods Alien and native aquatic plant species cover, water quality, forest canopy and adjacent land use were measured in three catchment locations (low‐, mid‐ and upper‐catchment) in all cities. Mean richness and cover of native and alien macrophytes were compared in the five cities, three catchment locations, and three land‐use types. Correlation tests examined relationships between alien and native richness at transect, site and city scales. Canonical correspondence analysis (CCA) determined the effects of environment on cover and richness of native and alien plant groups (emergents, floating, forbs/other, graminoids and submerged). Results Variation existed in the aquatic plant community at all scales, but strong patterns emerged with respect to land use and environmental gradients. Alien abundance was more responsive to anthropogenic disturbance (e.g. greater in intensive and agricultural land‐use types, and greater where dissolved nutrients and conductivity were high) than natives, which were unaffected by land‐use type and less responsive overall to environmental gradients. Native and alien richness were uncorrelated at all scales. Main conclusions Natives and aliens of the same life form did not respond similarly to the environment, suggesting inherent differences in their ability to capitalize on anthropogenic disturbance. Our results suggest invasion‐susceptible habitats are those that receive nutrient pollutants and that occur in urban and agricultural areas low in the catchment. Our confidence in these patterns is strengthened by their consistency across a large latitudinal gradient.     

Quantifying levels of biological invasion: towards the objective classification of invaded and invasible ecosystems

Biological invasions are a global phenomenon that threatens biodiversity, and few, if any, ecosystems are free from alien species. The outcome of human‐mediated introductions is affected by the invasiveness of species and invasibility of ecosystems, but research has primarily focused on defining, characterizing and identifying invasive species; ecosystem invasibility has received much less attention. A prerequisite for characterizing invasibility is the ability to compare levels of invasion across ecosystems. In this paper, we aim to identify the best way to quantify the level of invasion by nonnative animals and plants by reviewing the advantages and disadvantages of different metrics. We explore how interpretation and choice of these measures can depend on the objective of a study or management intervention. Based on our review, we recommend two invasion indices and illustrate their use by applying them to two case studies. Relative alien species richness and relative alien species abundance indicate the contribution that alien species make to a community. They are easy to measure, can be applied to various taxa, are independent of scale and are comparable across regions and ecosystems, and historical data are often available. The relationship between relative alien richness and abundance can indicate the presence of dominant alien species and the trajectory of invasion over time, and can highlight ecosystems and sites that are heavily invaded or especially susceptible to invasion. Splitting species into functional groups and examining invasion patterns of transformer species may be particularly instructive for gauging effects of alien invasion on ecosystem structure and function. Establishing standard, transparent ways to define and quantify invasion level will facilitate meaningful comparisons among studies, ecosystem types and regions. It is essential for progress in ecology and will help guide ecosystem restoration and management.     

Biotic resistance to plant invasion in grassland: Does seed predation increase with resident plant diversity?

Seed predation impacts heavily on plant populations and community composition in grasslands. In particular, generalist seed predators may contribute to biotic resistance, i.e. the ability of resident species in a community to reduce the success of non-indigenous plant invaders. However, little is known of predators’ preferences for seeds of indigenous or non-indigenous plant species or how seed predation varies across communities. We hypothesize that seed predation does not differ between indigenous and non-indigenous plant species and that seed predation is positively related to plant species diversity in the resident community. The seed removal of 36 indigenous and non-indigenous grassland species in seven extensively or intensively managed hay meadows across Switzerland covering a species-richness gradient of 18–50 plant species per unit area (c. 2 m²) was studied. In mid-summer 2011, c. 24,000 seeds were exposed to predators in Petri dishes filled with sterilized soil, and the proportions of seeds removed were determined after three days’ exposure. These proportions varied among species (9.2–62.5%) and hay meadows (17.8–48.6%). Seed removal was not related to seed size. Moreover, it did not differ between indigenous and non-indigenous species, suggesting that mainly generalist seed predators were active. However, seed predation was positively related to plant species richness across a gradient in the range of 18–38 species per unit area, representing common hay meadows in Switzerland. Our results suggest that generalist post-dispersal seed predation contributes to biotic resistance and may act as a filter to plant invasion by reducing the propagule pressure of non-local plant species.     

Defining fitness in evolutionary models

The analysis of evolutionary models requires an appropriate definition for fitness. In this paper, I review such definitions in relation to the five major dimensions by which models may be described, namely (i) finite versus infinite (or very large) population size, (ii) type of environment (constant, fixed length, temporally stochastic, temporally predictable, spatially stochastic, spatially predictable and social environment), (iii) density-independent or density-dependent, (iv) inherent population dynamics (equilibrium, cyclical and chaotic), and (v) frequency-dependent or independent. In simple models, the Malthusian parameter ‘r’ or the net reproductive rate R ₀ may be satisfactory, but once density-dependence or complex population dynamics is introduced the invasion exponent should be used. Defining fitness in a social environment or when there is frequency-dependence requires special consideration.     

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

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

不同光照强度对入侵植物薇甘菊（Mikania micrantha）和飞机草（Chromolaena odorata）生长及生物量分配的影响

外来植物入侵严重威胁着当地生物多样性和生态系统功能,引发巨大经济损失,已成为日益严重的全球性问题。光是热带和亚热带森林生态系统植物生长最重要的限制性资源之一,对光的捕获和利用直接影响植物在自然生态系统中的生存和适合度。本研究以华南地区危害严重的入侵植物薇甘菊(Mikania micrantha)和飞机草(Chromolaena odorata)为研究对象,对比二者生长、生物量分配及叶片特征对光照强度变化的响应,从而验证假设:外来植物入侵力受有效光辐射的限制,光照是影响不同演替阶段林型具有不同可入侵性的重要原因之一。研究结果表明,3%—60%全光照下两种外来植物均可存活,并显示较灵活的生物量分配策略,可见其对光照具有广适性,为其广泛分布提供生态学基础。低光严重抑制了两种外来植物的生物量积累,这可能是导致演替后期季风林入侵现象少的主要原因之一。对比两种植物叶片特征对光照强度变化的响应,高光下薇甘菊SLA降低,LAR、LARMR减小,叶片变小变厚,有利于植株保持强光下的水分平衡;低光下薇甘菊增加SLA和LAR,叶片变薄变大,增大对光的截获面积,以实现对光强减弱的适应。相比薇甘菊,3种光照处理对飞机草的SLA、LAR、LARMR无显著影响。以上结果提示:(1)光照影响所在群落对外来植物入侵的抵抗力;(2)攀援植物薇甘菊较丛生型草本飞机草对变化的光环境在叶片水平上具有更高的形态可塑性,有效寻觅光资源。     

Widespread resistance of Mediterranean island ecosystems to the establishment of three alien species

Although some invasive plants are cosmopolitan, not all ecosystems are invaded to the same degree. Yet there is little experimental work on how ecosystem resistance to invasion at the establishment phase differs among ecosystems. We conducted two field sowing experiments in two consecutive years to examine establishment of the deciduous tree Ailanthus altissima, the succulent subshrub Carpobrotus spp. and the annual geophyte Oxalis pes‐caprae in coastal dunes, shrublands and oldfields in more than 200 sites across six Mediterranean Basin islands differing in climatic conditions and local species richness. Establishment success (i.e. percentage of plots with at least one seedling) and rates (i.e. seedling to sown seed ratio) were low, especially for Ailanthus even when accounting for differences in seed viability. Oxalis was capable of producing a new cohort of seedlings the year following planting. By contrast, all Ailanthus seedlings and half the Carpobrotus seedlings died following the first summer. Differences in establishment success and rates among ecosystems were species‐, island‐ and year‐dependent. Differences in precipitation and mean temperature were associated with differences in establishment rates across sites. Establishment rates tended to be positively correlated with cumulative precipitation and negatively with mean T^a. Unexpectedly, native species richness was not a good predictor of seedling establishment, except for higher Oxalis establishment success in species rich habitats. By conducting field sowing tests at multiple sites across a region we found that except for Oxalis, Mediterranean island ecosystems are quite resistant to invader establishment. These results suggest that differences in the degree of invasion between ecosystems and islands might be more dependent upon the influence of invasion event factors (e.g. propagule pressure) or factors acting at a later life‐history stages rather than differences in the resistance imposed by ecosystems to invader recruitment. Moreover, our results support the notion that in Mediterranean ecosystems invasions are highly idiosyncratic events and strongly dependent on water availability conditions.     

Effects of experimental manipulation of light and nutrients on establishment of seedlings of native and invasive woody species in Long Island,NY forests

While earlier studies on the process of invasion often focused on single factors or on the general explanation of ‘disturbance,’ recent work has attempted to move towards a more mechanistic understanding of the factors that promote plant community invasion. Manipulative experiments provide a means for discerning causal relationships and interactive effects of environmental factors in promoting invasion; such experiments have been conducted in a number of grassland and shrub ecosystems. This study extends multifactor manipulative experiments into forest communities to compare factors influencing early seedling establishment for native and invasive woody plants. In Long Island, NY, invasion patterns are correlated with forest community type (pine barrens or hardwood), light availability, and soil N and Ca. We conducted manipulative field experiments in two different years to determine the relative importance and interaction of experimental gaps and N and Ca addition in pine barrens and hardwood forests in promoting invasion. We used seedlings of seven common native and invasive species in the first experiment, and 16 native and invasive species paired phylogenetically in the second experiment. Light had the strongest effect on plant growth; all plants grew more in gaps. We found no difference in the average growth rates of native and invasive species. Invasives responded more to high resources than did natives, with highest relative growth rates in gaps in the more fertile soils of the hardwood forests. Opportunities for invasion may differ from year to year, with differential success of invaders only in some years and under some environmental conditions. Clearly, to understand the complex interactions between resources and invasion in forests will require many manipulative experiments across a range of environments and using suites of invasive and native species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.