Evaluating differences in the shape of native and alien plant trait distributions will bring new insights into invasions of plant communities

Failure to quantify differences in the shape of inter‐specific trait distributions (e.g., skew, kurtosis) when comparing co‐occurring alien and native plants hinders the integration of biological invasions and plant community ecology. Within a plant community, understanding the circumstances that lead to the shape of the inter‐specific distribution of one or more functional plant traits being unimodal, bimodal, multimodal or skewed has the potential to shed new light on community vulnerability to invasion, subsequent ecosystem impacts and the selection pressures (e.g., stabilizing, directional or disruptive) acting upon native and alien species. Ignoring differences in the shape of inter‐specific trait distributions of alien and native species could miss important insights into plant invasions, including: the existence of unsaturated native plant communities, empty niches, shifting trait optima of species as a result of environmental change and incomplete colonization–extinction processes following invasion. Future comparisons of functional trait differences between native and alien species should include assessment of the shapes of inter‐specific trait distributions since these may differ even when the mean values of traits are similar for native and alien species. The infrequent application of such approaches may explain the limited generalizations regarding the drivers and consequences of plant invasions in plant communities.     

Release from belowground enemies and shifts in root traits as interrelated drivers of alien plant invasion success: a hypothesis

Our understanding of the interrelated mechanisms driving plant invasions, such as the interplay between enemy release and resource‐acquisition traits, is biased by an aboveground perspective. To address this bias, I hypothesize that plant release from belowground enemies (especially fungal pathogens) will give invasive plant species a fitness advantage in the alien range, via shifts in root traits (e.g., increased specific root length and branching intensity) that increase resource uptake and competitive ability compared to native species in the alien range, and compared to plants of the invader in its native range. Such root‐trait changes could be ecological or evolutionary in nature. I explain how shifts in root traits could occur as a consequence of enemy release and contribute to invasion success of alien plants, and how they could be interrelated with other potential belowground drivers of invasion success (allelopathy, mutualist enhancement). Finally, I outline the approaches that could be taken to test whether belowground enemy release results in increased competitive ability and nutrient uptake by invasive alien plants, via changes in root traits in the alien range.     

Invasive plants in Minnesota are “joining the locals”: A trait‐based analysis

Questions

Predicting which newly arrived species will establish and become invasive is a problem that has long vexed researchers. In a study of cold temperate oak forest stands, we examined two contrasting hypotheses regarding plant functional traits to explain the success of certain non‐native species. Under the “join the locals” hypothesis, successful invaders are expected to share traits with resident species because they employ successful growth strategies under light‐limited understorey conditions. Instead, under the “try harder” hypothesis, successful invaders are expected to have traits different from native species in order to take advantage of unused niche space.

Location

Minnesota, USA.

Methods

We examined these two theories using 109 native and 11 non‐native plants in 68 oak forest stands. We focused on traits related to plant establishment and growth, including specific leaf area (SLA), leaf carbon‐to‐nitrogen ratio (C:N), wood density, plant maximum height, mycorrhizal type, seed mass and growth form. We compared traits of native and non‐native species using ordinations in multidimensional trait space and compared community‐weighted mean (CWM) trait values across sites.

Results

We found few differences between trait spaces occupied by native and non‐native species. Non‐native species occupied smaller areas of trait space than natives, yet were within that of the native species, indicating similar growth strategies. We observed a higher proportion of non‐native species in sites with higher native woody species CWM SLA and lower CWM C:N. Higher woody CWM SLA was observed in sites with higher soil pH, while lower CWM C:N was found in sites with higher light levels.

Conclusions

Non‐native plants in this system have functional traits similar to natives and are therefore “joining the locals.” However, non‐native plants may possess traits toward the acquisitive end of the native plant trait range, as evidenced by higher non‐native plant abundance in high‐resource environments.

     

Responses of macrophytes to dewatering: effects of phylogeny and phenotypic plasticity on species performance

Temporary dewatering constitutes a drastic change in conditions for aquatic vegetation. Species’ sustained performance under these conditions relies partly on their ability to produce a terrestrial phenotype. Such adaptations may include the development of self-supporting aboveground organs with higher dry matter content enabling plants to withstand gravity and smaller leaves with thicker cuticle to reduce evapotranspiration, leading to lower specific leaf area, higher leaf-construction costs and consequently higher leaf life span. The ability of aquatic plant species to produce a terrestrial-adapted phenotype may differ according to growth form and evolutionary history. The objectives of this study were to (1) measure the effects of dewatering on aquatic plant performance, (2) determine how growth form and phylogenetic position affect performance, and (3) relate plant performance to plasticity. To meet these objectives, we experimentally studied aquatic plant responses to dewatering by measuring survival, growth, and a set of traits describing the morphology and leaf-resource economy of eight aquatic plant species with contrasting phylogeny and growth forms. The ability of aquatic plants to withstand dewatering differed according to phylogeny but not to growth form. The eudicots presented high survival and similar growth rates under terrestrial compared to aquatic conditions, while monocots generally did not survive dewatering. These species produced phenotypic adjustments, such as denser aboveground organs and leaf plasticity, which can explain the maintenance of similar growth rates under terrestrial conditions. The relatively strong plasticity and performance of eudicots in terrestrial habitats suggests that their optimal niche is the interface between aquatic and terrestrial ecosystems.     

Do alien and native tree species from Central Argentina differ in their water transport strategy?

In this study we assessed the water transport strategies and the abundance of alien and native tree species at a regional scale in Córdoba Mountains, Central Argentina. The aims of this study were: (i) to analyse whether alien and native tree species show divergent water transport strategies; and (ii) to explore whether species abundances of alien and natives are associated with specific trait attributes. Eight alien species and 12 native species were recorded in 50 complete vascular plant vegetation relevés. Water transport strategies were assessed through the following functional traits: minimum leaf water potential, potential water content of wood, effective leaf area, leaf area per sapwood area and wood density. Also, resource use strategies were assessed throughout the measurement of specific leaf area. We found that alien species had a higher efficiency in water transport (i.e. higher minimum leaf water potential and lower wood density values) and faster resource acquisition and use (higher specific leaf area values) than native species. We did not find evidence suggesting that the relative abundance of species was associated to water transport strategies and faster resource acquisition and use. Alien species seem to differ from natives in specific functional attributes that are absent in the resident community and might allow aliens to use more resources and at a higher rate than native species. Finally, our results show the potential of a trait‐based predictive framework for alien species, and the possible effects on ecosystem functions.     

Climate change and the invasion of California by grasses

Over the next century, changes in the global climate are expected to have major consequences for plant communities, possibly including the exacerbation of species invasions. We evaluated this possibility in the grass flora of California, which is economically and ecologically important and heavily invaded. We used a novel, trait‐based approach involving two components: identifying differences in trait composition between native and exotic components of the grass flora and evaluating contemporary trait–climate relationships across the state. The combination of trait–climate relationships and trait differences between groups allows us to predict changes in the exotic‐native balance under climate change scenarios. Exotic species are more likely to be annual, taller, with larger leaves, larger seeds, higher specific leaf area, and higher leaf N percentage than native species. Across the state, all these traits are associated with regions with higher temperature. Therefore, we predict that increasing temperatures will favor trait states that tend to be possessed by exotic species, increasing the dominance of exotic species. This prediction is corroborated by the current distribution of exotic species richness relative to native richness in California; warmer areas contain higher proportions of exotic species. This pattern was very well captured by a simple model that predicts invasion severity given only the trait–climate relationship for native species and trait differences between native and exotic species. This study provides some of the first evidence for an important interaction between climate change and species invasions across very broad geographic and taxonomic scales.     

Impact of invasions by alien plants on soil seed bank communities: Emerging patterns

Much of our current understanding of the impact of invasive species on plant communities is based on patterns occurring in the above-ground vegetation, while only few studies have examined changes in soil seed banks associated with plant invasions, despite their important role as determinants of vegetation dynamics. Here, we reviewed the literature on the impact of plant invasions on the seed bank and we provide a quantitative synthesis using a meta-analysis approach. Specifically, (1) we quantified the impact of 18 invasive alien plants on (i) species richness and (ii) density of the seed banks of invaded communities, based on 58 pair-wise invaded-uninvaded comparisons (cases); we identified (2) the invasive taxa that are responsible for the largest changes in the seed bank; and (3) the habitats where substantial changes occur. Our study showed three major findings: (1) species richness (68% of cases) and density (58% of cases) were significantly lower in native seed banks invaded by alien plants; (2) species richness and density of native and alien species were remarkably lower in seed banks invaded by large, perennial herbs compared to uninvaded sites; and (3) invaded seed banks were often associated with a larger richness and/or abundance of alien species. This study indicates a need for additional seed bank data in invasion ecology to characterize species-specific and habitat-specific impacts of plant invasions, and to determine whether changes in the seed banks of native and alien species are a symptom of environmental degradation prior to a plant invasion or whether they are its direct result. The findings of this study help improve our capacity to predict the long-term implications of plant invasions, including limitations in the recruitment of native species from the seed bank and the potential for secondary invasions by seeds of other alien species.     

From introduction to nuisance growth: a review of traits of alien aquatic plants which contribute to their invasiveness

Invasive alien aquatic plant species (IAAPs) cause serious ecological and economic impact and are a major driver of changes in aquatic plant communities. Their invasive success is influenced by both abiotic and biotic factors. Here, we summarize the existing knowledge on the biology of 21 IAAPs (four free-floating species, eight sediment-rooted, emerged or floating-leaved species, and nine sediment-rooted, submerged species) to highlight traits that are linked to their invasive success. We focus on those traits which were documented as closely linked to plant invasions, including dispersal and growth patterns, allelopathy and herbivore defence. The traits are generally specific to the different growth forms of IAAPs. In general, the species show effective dispersal and spread mechanisms, even though sexual and vegetative spread differs strongly between species. Moreover, IAAPs show varying strategies to cope with the environment. The presented overview of traits of IAAPs will help to identify potential invasive alien aquatic plants. Further, the information provided is of interest for developing species-specific management strategies and effective prevention measures.

     

Are traits measured on pot grown plants representative of those in natural communities?

Question: The quantification of functional traits in natural communities can be difficult (e.g. root traits, RGR). Can functional traits measured on pot grown plants be reliably applied to natural communities? Alternatively, can below‐ground plant traits be predicted from above‐ground traits? Location: Southeastern Australia. Methods: We compared 17 shoot, root and whole‐plant morphological traits measured on 14 plant species in a native grassland community to those measured under two different pot conditions: unfertilised and fertilised. Results: The majority of trait values for pot grown plants differed to plants in the field, however, species ranking remained consistent for most leaf traits between the field and the two pot growing conditions. In contrast, species ranking was not consistent for most whole plant traits when comparing field plants to fertilised pot grown plants, providing a caution against the tendency to grow plants in controlled conditions at ‘optimal’ (high) resource levels. Moderate to strong correlations were found between below‐ground and above‐ground plant traits, including between root dry matter content and leaf dry matter content, and between specific root area and specific leaf area. Conclusions: The utility of pot grown plants to quantify traits for field plants is highly dependent on the selection of the growing conditions in the controlled environment. The consistency we observed between above‐ground and below‐ground trait strategies suggests that below‐ground traits may be predictable based on above‐ground traits, reducing the need to quantify root traits on cultured plants.     

Invasive plants do not display greater phenotypic plasticity than their native or non‐invasive counterparts: a meta‐analysis

Phenotypic plasticity is commonly considered as a trait associated with invasiveness in alien plants because it may enhance the ability of plants to occupy a wide range of environments. Although the evidence of greater phenotypic plasticity in invasive plants is considerable, it is not yet conclusive. We used a meta‐analysis approach to evaluate whether invasive plant species show greater phenotypic plasticity than their native or non‐invasive counterparts. The outcome of such interspecific comparisons may be biased when phylogenetic relatedness is not taken into account. Consequently, species pairs belonged to the same genus, tribe or family. The meta‐analysis included 93 records from 35 studies reporting plastic responses to light, nutrients, water, CO₂, herbivory and support availability. Contrary to what is often assumed, overall, phenotypic plasticity was similar between invasive plants and native or non‐invasive closely related species. The same result was found when separate analyses were conducted for trait plasticity to nutrients, light and water availability. Thus, invasive plant species and their native or non‐invasive counterparts are equally capable of displaying functional responses to environmental heterogeneity. The colonization of a wide range of environments by invasive plants could be due to their capacity to undergo adaptive ecotypic differentiation rather than to their ability to display plastic responses. Alternatively, phenotypic plasticity might play a role in plant invasion, but only during the initial phases, when tolerance of the novel environment is essential for plant survival. Afterwards, once alien plants are identified as invaders, the magnitude of phenotypic plasticity might be reduced after selection of the optimum phenotypes in each habitat. The identification of plant traits that consistently predict invasiveness might be a futile task because different traits favor invasiveness in different environments. Approaches at the local scale, focusing on the ecology of specific invasive plants, could be more fruitful than global macro‐analyses.     

潮虫消耗木本植物凋落物的可选择性试验

近年来,越来越多的学者关注外来植物入侵对土壤生态功能的影响效应及其相应反馈机制的探索与研究,然而本地原生土壤生物群落对不同入侵程度下的外来植物种以及本地原生植物种之间是否存在消耗差异却尚不明了.通过等足目潮虫的选择性喂养试验来测试10个本地种、5个非入侵性外来种和5个强入侵性外来种之间的适口性差异,试图求证外来植物的入侵性是否与植物落叶被消耗率呈现必然联系.数据分析结果显示潮虫对本地种、非入侵性外来种和入侵性外来种的消耗并无显著差异;而潮虫对不同生活型下木本植物的取食却存在显著差异,即灌木消耗率显著高于乔木.其次,通过植物初始性状指标(包括木质素、纤维素、半纤维素、碳、氮含量)与相应消耗率的相关比较,消耗量总体上与植物凋落物的氮含量呈正相关关系(R2 =0.358).由此,研究结论强调植物落叶的降解速率并不一定与植物入侵性或入侵阶段呈绝对相互关连,但是氮含量,抑或各种形式的植物氮元素成分都可能在一定程度上参与并影响着外来植物的入侵进程.     

Phenotypic trait differences between Iris pseudacorus in native and introduced ranges support greater capacity of invasive populations to withstand sea level rise

Aim

Tidal wetlands are greatly impacted by climate change, and by the invasion of alien plant species that are being exposed to salinity changes and longer inundation periods resulting from sea level rise. To explore the capacity for the invasion of Iris pseudacorus to persist with sea level rise, we initiated an intercontinental study along estuarine gradients in the invaded North American range and the native European range.

Location

San Francisco Bay-Delta Estuary; California, USA and Guadalquivir River Estuary; Andalusia, Spain.

Methods

We compared 15 morphological, biochemical, and reproductive plant traits within populations in both ranges to determine if specific functional traits can predict invasion success and if environmental factors explain observed phenotypic differences.

Results

Alien I. pseudacorus plants in the introduced range had more robust growth than plants in the native range. The vigour of the alien plants was reflected by expression of higher leaf water content, fewer senescent leaves per leaf fan, and more carbohydrate storage reserves in rhizomes than plants in the native range. Moreover, alien plants tended to show higher specific leaf area and seed production than native plants. I. pseudacorus plants in the introduced range were less affected by increasing salinity and were exposed to deeper inundation water along the estuarine gradient than those in the native range.

Main Conclusions

Functional trait differences suggest mature populations of I. pseudacorus in the introduced range have greater adapted capacity to adjust to environmental stresses induced by rising sea level than those in the native range. Knowledge of these trait responses can be applied to improve risk assessments in invaded estuaries and to achieve climate-adapted conservation goals for conservation of the species in its native range.     

A new comprehensive database of alien plant species in Chile based on herbarium records

There is an urgent need for comprehensive national databases on alien plant species, especially in developing countries. Despite the fact that plant invasions are considered a major threat to biodiversity, they have been poorly studied or not considered a conservation priority in South America. We aim to assess alien plant distribution in Chile, using the first comprehensive public alien plant database, and discuss the implications of using herbarium records to develop national databases of alien plants. We used herbarium records to assemble a comprehensive national database of alien plants. We calculated the number of alien and native species and specimens recorded in each 10 × 10 km cell. We evaluated sampling efforts and tested for relationships between alien and native species collections, as well as other spatial patterns along the latitudinal gradient. Alien and native species richness was positively correlated. Alien plants were mostly collected in central Chile, with few species collected in both the extreme north and south. However, native plants were strongly collected in central Chile, as well as in both extremes of the country. Alien and native plants followed the same pattern of accumulation along the latitudinal gradient, with native plants being relatively more collected than alien plants. Herbarium records provide valuable baseline information to evaluate plant species distribution. However, there are important gaps in this database, (e.g. variable sampling effort for alien and native plants, incomplete information on life-history traits). Given scientists and land managers increasing demand for baseline information and the high cost of collecting such data in developing countries, herbarium records should be used more frequently for research and management of plant invasions.     

植物功能性状与外来植物入侵

揭示影响外来植物入侵性的功能性状及其生态机制是入侵植物生态学的核心任务之一。本文综述了植物功能性状与外来植物入侵性的研究进展, 通过分析植物功能性状对外来植物入侵的贡献以及外来植物的不同入侵阶段对其功能性状的需求, 探讨植物功能性状与外来植物入侵的相关性及其入侵机理。迄今研究较多的影响外来植物入侵性的功能性状主要包括形态性状、生长性状、生理性状、繁殖性状、种子性状、克隆性状、表型可塑性和遗传变异等。这些功能性状对外来植物入侵的贡献随着入侵阶段的不同而变化。在传播到达阶段, 种子性状对入侵具有重要影响; 在定居建群阶段, 与植物抗逆性和适应性相关的生理性状和繁殖性状发挥主要作用; 在扩散入侵阶段, 克隆性状和影响植物竞争能力的生理性状对植物成功入侵具有重要贡献。由于植物入侵性是其功能性状和环境因素互作的结果, 且功能性状的作用随环境因素和入侵阶段不同而异, 因此, 结合外来植物入侵阶段, 并考虑功能性状与环境因子的互作, 是入侵生物学中植物功能性状研究的发展趋势。     

Faster returns on ‘leaf economics’ and different biogeochemical niche in invasive compared with native plant species

Plant‐invasive success is one of the most important current global changes in the biosphere. To understand which factors explain such success, we compared the foliar traits of 41 native and 47 alien‐invasive plant species in Oahu Island (Hawaii), a location with a highly endemic flora that has evolved in isolation and is currently vulnerable to invasions by exotic plant species. Foliar traits, which in most cases presented significant phylogenetic signal, i.e. closely related species tended to resemble each other due to shared ancestry, separated invasive from native species. Invasive species had lower leaf mass per area and enhanced capacities in terms of productivity (photosynthetic capacity) and nutrient capture both of macro‐ (N, P, K) and microelements (Fe, Ni, Cu and Zn). All these differences remain highly significant after removing the effects of phylogenetic history. Alien‐invasive species did not show higher efficiency at using limiting nutrient resources, but they got faster leaf economics returns and occupied a different biogeochemical niche, which helps to explain the success of invasive plants and suggests that potential increases in soil nutrient availability might favor further invasive plant success.     

Plants for nitrogen management in riparian zones: A proposed trait‐based framework to select effective species

Restoring plants to the riparian zone is regarded as management best practice in river restoration and has the potential to reduce the impact of nitrogen (N) pollution on aquatic organisms and improve water quality for human use. Plant characteristics and the interplay of hydrology and biogeochemistry control N retention in the riparian zone. The balance between processes such as denitrification and plant assimilation determines riparian N retention. Plant traits are likely to mediate these N removal processes through variations in root form, growth character, foliage production (quantity, quality and rate of return to the soil) and by altering conditions in the rhizosphere soil. Vegetation can slow N transfer via direct plant uptake of N (during periods of rapid vegetation growth) and changes induced to soil hydrology, nutrient cycling and microbial activity, principally denitrification. Few studies have focused on species‐dependent effects on N movement through soil and across boundaries. We propose a new framework, based on a literature review of plant traits with respect to N cycling, which can be used to select plant species with traits likely to maximise N removal during transport through the riparian zone. In the proposed framework, inter‐specific differences in traits known to influence N mobility: root form, growth rate, foliar characteristics and rhizosphere processes, are used to describe species’ potential impact on N removal. Plant trait data may be drawn from studies outside the riparian zone; for example forest ecology, horticulture or forestry research, and candidate species are scored to predict N removal efficiency. We apply the framework to New Zealand's native riparian plant assemblages to demonstrate the trait‐based approach. This framework can guide restoration management decisions and investment in riparian revegetation in a manner that is not restricted to geographically specific or well‐studied species.     

Biological invasion hotspots: a trait‐based perspective reveals new sub‐continental patterns

Invader traits (including plant growth form) may play an important, and perhaps overlooked, role in determining macroscale patterns of biological invasions and therefore warrant greater consideration in future investigations aimed at understanding these patterns. To assess this need, we used empirical data from a national‐level survey of forest in the contiguous 48 states of the USA to identify geographic hotspots of forest plant invasion for three distinct invasion characteristics: invasive species richness, trait richness (defined as the number of the five following plant growth forms represented by the invasive plants present at a given location: forbs, grasses, shrubs, trees, and vines), and species richness within each growth form. Three key findings emerged. 1) The hotspots identified encompassed from 9 to 23% of the total area of our study region, thereby revealing many forests to be not only invaded, but highly invaded. 2) Substantial spatial disagreement among hotspots of invasive species richness, invasive trait richness, and species richness of invasive plants within each growth form revealed many locations to be hotspots for invader traits, or for particular growth forms of invasive plants, rather than for invasive plants in general. 3) Despite eastern forests exhibiting higher levels of plant invasion than western forests, species richness for invasive forbs and grasses in the west were respectively greater than and equivalent to levels found in the east. Contrasting patterns between eastern and western forests in the number of invasive species detected for each growth form combined with the spatial disagreement found among hotspot types suggests trait‐based variability in invasion drivers. Our findings reveal invader traits to be an important contributor to macroscale invasion patterns.     

Root morphological and structural comparisons of introduced and native aquatic plant species in multiple substrates

Invasions of introduced plants are considered among the greatest threats to biodiversity worldwide. Aquatic habitats suffer invasion more frequently and extensively than do terrestrial habitats. Although the role of roots in plant anchoring and support is important, previous studies have focused much attention on the morphological traits of above-ground parts, with relatively less attention given to the root structures of aquatic plants. In this study, we aimed to compare differences in root morphological and structural traits between introduced and native plants in response to different substrates. We hypothesized that introduced aquatic plants have an advantage over native plants with regard to root trait values and plasticity. A total of six aquatic plants were used: Two invasive and one exotic noninvasive species were paired with their native counterparts according to life form (amphibious emergent, submerged and floating-leaved) and cultivated in substrates of clay, a clay/sand mixture (v:v = 1:1) or sand. Root morphological traits, topological indices and root relative distance plasticity indices were quantified. The results indicated that different substrates have various effects on the root traits of these six aquatic plants; the introduced plants generally exhibited higher plasticity than did their native counterparts of the same life form.     

Low resource availability limits weed invasion of tropical savannas

The savanna biome is one of the least invaded among global biomes, although the mechanisms underpinning its resistance to alien species relative to other biomes is not well understood. Invaders generally are at the resource acquisitive end of functional global plant trait variation and in low-resource savanna environments we might expect that successful invaders will only outperform native species under resource rich or highly disturbed conditions. However, invaders may also directly exploit resource stressed environments using resource conservative traits in some situations. It’s also possible that successful invaders and native species largely overlap in their trait profiles indicating site specific environmental factors are responsible for invader success in particular contexts rather than a general trait and functional divergence between invaders and native species. To address these various hypotheses, we compared a suite of morphological and physiological traits in graminoid and herbaceous native and co-occurring invasive plant species across a range of habitats in savannas of the Kimberley region of northern Australia. Invader grass species had traits associated with resource acquisition and fast growth rates, such as high SLA and leaf nutrient contents. In contrast, dominant native perennial grasses had traits characteristic of resource conservation and slow growth in resource stressed conditions. Trait profiles among invasive forbs and legumes exhibited stress tolerant traits relative to their native counterparts. Invaders also displayed strong divergence in reproductive traits, suggesting diverse responses to disturbance not indicated by leaf economic traits alone. These results suggest that savannas may be resistant to invaders with resource acquisitive traits due to their strong resource limitation.