Trait values and not invasive status determine competitive outcomes between native and invasive species under varying soil nutrient availability

Invasion by exotic plants is often associated with nutrient enrichment of soils, particularly on soils of naturally low fertility. As a consequence, it is likely that the outcome of competitive interactions between native and invasive plants may be mediated by soil nutrient availability. We independently investigated competitive effect and response as well as the occurrence of asymmetric competition among native and invasive plants on soils of varying nutrient availability, using a glasshouse experiment. Seedlings of eight co‐occurring pairs of invasive and native species from low fertility Hawkesbury Sandstone‐derived soil were grown under low and high nutrient availability. We tested the hypotheses that native species would be competitively superior at low nutrient availability and have trait values associated with a resource conservation strategy while invasive species would be competitively superior at high nutrient availability and have trait values associated with a resource acquisition strategy. We found that nutrient availability did not mediate competitive interactions between invasive and native species. Instead, two invasive and one native species were always competitively superior irrespective of nutrient availability. Competitively superior species displayed a mixture of both resource conservation and acquisition strategies at low and high nutrient availability. In support of previous studies, we found that the a priori classification of invasive and native species does not predict competitive superiority at varying nutrient levels. Rather, species specific differences in trait values provide a competitive advantage in response to nutrient availability.     

Nutrient stress and performance of invasive Hieracium lepidulum and co-occurring species in New Zealand

Many studies have highlighted the relationship between nutrient fluctuations and enrichment in the process of plant invasion. However, invasion may be also associated with conditions of plant stress, either nutrient depletion or toxicity, in the environment. In this study, we investigate the possible role of nutrient stress in the invasion of Hieracium lepidulum (Stenstroem) Omang, in South Island, New Zealand. We do this by comparing several performance attributes, and their plasticity, for H. lepidulum and a number of co-occurring species, across a series of nutrient depletion and toxicity tests. H. lepidulum had intermediate yields, high root:shoot ratios and high tissue nutrient contents at control nutrient concentrations. H. lepidulum differed in edaphic tolerance from all but Chionochloa flavescens var. brevis, in being insensitive to nutrient dilutions other than nitrogen. The significance of performance in terms of edaphic tolerance and adaptations are discussed. Intermediate yields at control nutrient levels suggest that H. lepidulum should not be competitive compared to high yielding species including Agrostis stolonifera and Poa cita, but should be competitive with lower yielding Coprosma rugosa and C. flavescens var. brevis. Conversely, significant yield decreases under nitrogen limitation stress suggests that H. lepidulum will not likely occupy very nutrient poor sites. H. lepidulum, along with C. flavescens var. brevis, were found to be tolerant of ammonium as an alternative source of nitrogen, while other species were not. These data suggest that H. lepidulum and C. flavescens var. brevis would be relatively tolerant of the stresses associated with acidic soils compared to the other species, but not to stresses associated with absolute shortage of nitrogen. Combined results point to the likely occurrence of H. lepidulum at sites of intermediate fertility. The possible roles of ammonium stress and disturbance reliance in further defining H. lepidulum ecology is discussed.     

Vulnerability of Mediterranean Basin ecosystems to climate change and invasion by exotic plant species

Aim To assess at a broad scale the vulnerability of Mediterranean vegetation to alien plant invasion under different climatic and disturbance scenarios. Location We simulated the vegetation biogeography and dynamics on five of the main islands of the Mediterranean Basin: Mallorca, Corsica, Sardinia, Crete and Lesvos. Methods We used LPJ‐GUESS, a generalized ecosystem model based on dynamic processes describing establishment, competition, mortality and ecosystem biogeochemistry. We simulated the vegetation distribution and dynamics using a set of plant functional types (PFTs) based on bioclimatic and physiological parameters, which included tree and shrub PFTs defined especially for the Mediterranean. Additionally, two invasive PFTs, an invasive tree type and an invasive herb type, were defined and used to estimate the vulnerability to invasion of a range of different ecosystems. The model was used to simulate climate changes and associated changes in atmospheric [CO₂] to 2050 according to two SpecialReport on Emissions Scenarios climate scenarios (A1Fi and B1) combined with mean disturbance intervals of 3 and 40 years. Results The simulations and scenarios showed that the effect of climate change alone is likely to be negligible in many of the simulated ecosystems, although not all. The simulated progression of an invasion was highly dependent on the initial ecosystem composition and local environmental conditions, with a particular contrast between drier and wetter parts of the Mediterranean, and between mountain and coastal areas. The rate of ecosystem disturbance was the main factor controlling susceptibility to invasion, strongly influencing vegetation development on the shorter time scale. Main conclusions Further invasion into Mediterranean island ecosystems is likely to be an increasing problem: our simulations predict that, in the longer term, almost all the ecosystems will be dominated by exotic plants irrespective of disturbance rates.     

Current velocity shapes co‐existence patterns among invasive Dikerogammarus species

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Variation in performance of two co‐occurring mosquito species across diverse resource environments: insights from nutrient and stable isotope analyses

1. Inputs of animal and plant detritus are the main energy sources for food webs in a number of isolated container systems, including discarded automobile tyres and tree holes. Containers are dominated by mosquitoes in the genera Culex and Aedes, which among other differences often engage in different foraging behaviours. We hypothesised that because Aedes feed more by browsing surfaces, whereas Culex often filter the water column, these mosquitoes would show variation in performance and differentially affect detritus. Effects of different ratios of animal and plant detritus on survival, mass, and development time for two common container mosquito species, Culex restuans L. and Aedes albopictus Skuse, were examined. We also quantified detrital contribution to biomass via isotopic and nutrient analysis and the effect of larvae on detrital decay. 2. Adult male and female mass of both species was highest with some animal detritus and lowest in only leaf detritus. Aedes albopictus survival was higher than C. restuans across most detritus ratios. 3. Aedes albopictus had higher values of ¹⁵N and in some cases ¹³C across all detritus ratios compared with C. restuans; A. albopictus had lower nitrogen in tissue. Aedes albopictus appeared to be more efficient at obtaining potentially limiting nutrients and had a greater overall effect on detrital decay – a possible consequence of greater foraging effort. 4. Findings further support the view that mosquito performance can be influenced by detritus type, and provide a more precise hypothesis (i.e. lower need for nitrogen) that may explain the superior competitive ability of A. albopictus over other container mosquitoes.     

Is there evidence for the post‐invasion evolution of increased size among invasive plant species?

Many plant species grow taller and have higher reproductive capacity where they are nonindigenous invaders than where they are native components of the flora. Traditionally, it has been accepted that this is a plastic response to a benign environment, though recently this assumption has been challenged and a genetic basis for increased plant size has been invoked. We tested the hypothesis that the increased size of certain weed species is genetically, rather than environmentally, based. A common environment growth experiment revealed no significant differences in the size of Carduus nutans , Digitalis purpurea , Echium vulgare or Senecio jacobaea sampled from alien (Australia and New Zealand) or native (Britain and continental Europe) habitats. We conclude that post-invasion genetic changes associated with increased size may be unusual and that the phenomenon, where it occurs, generally reflects a plastic response to a novel environment.     

Life‐history traits predict perennial species response to fire in a desert ecosystem

The Mojave Desert of North America has become fire‐prone in recent decades due to invasive annual grasses that fuel wildfires following years of high rainfall. Perennial species are poorly adapted to fire in this system, and post‐fire shifts in species composition have been substantial but variable across community types. To generalize across a range of conditions, we investigated whether simple life‐history traits could predict how species responded to fire. Further, we classified species into plant functional types (PFTs) based on combinations of life‐history traits and evaluated whether these groups exhibited a consistent fire‐response. Six life‐history traits varied significantly between burned and unburned areas in short (up to 4 years) or long‐term (up to 52 years) post‐fire datasets, including growth form, lifespan, seed size, seed dispersal, height, and leaf longevity. Forbs and grasses consistently increased in abundance after fire, while cacti were reduced and woody species exhibited a variable response. Woody species were classified into three PFTs based on combinations of life‐history traits. Species in Group 1 increased in abundance after fire and were characterized by short lifespans, small, wind‐dispersed seeds, low height, and deciduous leaves. Species in Group 2 were reduced by fire and distinguished from Group 1 by longer lifespans and evergreen leaves. Group 3 species, which also decreased after fire, were characterized by long lifespans, large non‐wind dispersed seeds, and taller heights. Our results show that PFTs based on life‐history traits can reliably predict the responses of most species to fire in the Mojave Desert. Dominant, long‐lived species of this region possess a combination of traits limiting their ability to recover, presenting a clear example of how a novel disturbance regime may shift selective environmental pressures to favor alternative life‐history strategies.     

Trait–performance relationships of grassland plant species differ between common garden and field conditions

The way functional traits affect growth of plant species may be highly context‐specific. We asked which combinations of trait values are advantageous under field conditions in managed grasslands as compared to conditions without competition and land‐use. In a two‐year field experiment, we recorded the performance of 93 species transplanted into German grassland communities differing in land‐use intensity and into a common garden, where species grew unaffected by land‐use under favorable conditions regarding soil, water, and space. The plants’ performance was characterized by two independent dimensions (relative growth rates (RGR) of height and leaf length vs. aboveground biomass and survival) that were differently related to the eight focal key traits in our study (leaf dry matter content (LDMC), specific leaf area (SLA), height, leaf anatomy, leaf persistence, leaf distribution, vegetative reproduction, and physical defense). We applied multivariate procrustes analyses to test for the correspondence of the optimal trait–performance relationships between field and common garden conditions. RGRs were species‐specific and species ranks of RGRs in the field, and the common garden were significantly correlated. Different traits explained the performance in the field and the common garden; for example, leaf anatomy traits explained species performance only in the field, whereas plant height was found to be only important in the common garden. The ability to reproduce vegetatively, having leaves that are summer‐persistent and with high leaf dry matter content (LDMC) were traits of major importance under both settings, albeit the magnitude of their influence differed slightly between the field and the common garden experiment. All optimal models included interactions between traits, pointing out the necessity to analyze traits in combination. The differences between field and common garden clearly demonstrate context dependency of trait‐based growth models, which results in limited transferability of favorable trait combinations between different environmental settings.     

Geographical distributions in tropical trees: can geographical range predict performance and habitat association in co‐occurring tree species?

Aim A major floristic and climatic transition from aseasonal to seasonal evergreen tropical forest (the Kangar–Pattani Line; KPL) exists in the Indo‐Sundaic region of Southeast Asia. Mechanisms constraining species distribution here are at present poorly understood, but it is hypothesized that species differ in their tolerances of abiotic factors, in particular water availability. Under this hypothesis, we anticipate differences in performance or habitat preferences, or both, of species differing in distribution with respect to the KPL. The aim of this study is to test whether geographical distributions can be used to explain variation in growth, mortality and habitat preferences in co‐occurring tree species differing in their distribution in relation to the KPL. Location Pasoh Forest Reserve, Negeri Sembilan, Malaysia; south of the KPL. Methods All tree species within a 50‐ha forest dynamics plot were classified as widespread or southern based upon their distributions in relation to the KPL and as habitat specialists or generalists based on spatial association with soil‐based habitat categories. Growth and mortality rates, variation in growth and mortality with respect to soil type, and levels of habitat association were quantified for species with different geographical distributions. Results Differences existed in species performance based upon geographical distributions. Specifically, widespread species had lower growth rates than did species restricted to the aseasonal forests. Mortality rates did not differ as a function of geographical distribution. The growth responses of species to soil habitats also diverged, such that differences in performance of widespread species among soil types were more conservative than those of species restricted in their distribution to the aseasonal forests. However, the proportion of species showing positive habitat associations did not differ significantly between widespread and southern species. Main conclusions Distribution‐based differences in species performance and response to soil type support the hypothesis that species tolerant of wider climatic variation perform less well in any given environment due to limitations on plasticity. These performance differences provide quantitative evidence of the role of climatic transitions in determining tree species distributions in relation to the Kangar–Pattani Line in the Indo‐Malay region. Such differences in performance have important implications for our understanding of biodiversity gradients and responses of Indo‐Sundaic forests to climate change.     

Examining the link between competition and negative co‐occurrence patterns

Negative species co‐occurrence patterns have long intrigued ecologists because of their potential link to competition. Although manipulative field experiments have consistently revealed evidence of competition in natural communities, there is little evidence that this competition produces negative co‐occurrence patterns. Evidence does suggest that abiotic variation, dispersal limitation and herbivory can contribute to patterns of negative co‐occurrence among species; it is possible these influences have obscured a link with competition. Here, we test for a connection between negative co‐occurrence and competition by examining a small‐scale, relatively homogeneous old‐field plant community where the influence of abiotic variation was likely to be minimal and we accounted for the impact of herbivory with an herbivore exclosure treatment. Using three years of data (two biennial periods), we tested whether negatively co‐occurring pairs of species, when occasionally found together, experienced asymmetric abundance decline more frequently than positively co‐occurring pairs, for which there is no such expectation. We found no evidence that negatively co‐occurring pairs consistently suffered asymmetric abundance decline more frequently than positively co‐occurring pairs, providing no evidence that competition is a primary driver of negative co‐occurrence patterns in this community. Our results were consistent across control and herbivore exclosure treatments, suggesting that herbivores are not driving patterns of negative species co‐occurrence in this community. Any influence of competition or herbivory on co‐occurrence patterns is small enough that it is obscured by other factors such as substrate heterogeneity, dispersal and differential species responses to climatic variation through time. We interpret our results as providing evidence that competition is not responsible for producing negative co‐occurrence patterns in our study community and suggest that this may be the case more broadly.     

Local performance of six clonal alien species differs between native and invasive regions in Germany and New Zealand

Exotic plant invasions are widely observed to have strong biogeographic patterns with invasive species occurring at higher abundances in their introduced range when compared with their native range. However, only few field studies have validated this assumption by comparing plant populations of multiple species in their native and introduced ranges and have evaluated to what extent changes in sexual and clonal reproduction potentially have contributed to the success of plant invasions. Here, we present the results of a comparative field study in both the native (Germany) and the introduced (New Zealand, NZ) ranges of six clonal plant species with different invasive status: Achillea millefolium L., Pilosella officinarum Vaill., Hypericum perforatum L., Prunella vulgaris L., Leucanthemum vulgare Lam. and Lotus pedunculatus Cav. We hypothesized that all six species show better performance in introduced NZ than in native German populations and tested if population structures investigated at different scales provide a useful tool to identify differences between native and introduced occurrences. In 10 populations per species and country we assessed plant density and flowering proportion at the population scale and around individual plants, thereby identifying the ‘crowdedness’ of the populations. Furthermore, we collected individual plants and determined the number of attached clonal organs and plant biomass. For all six species crowdedness in NZ populations was higher than in German populations. Additionally, overall population density of four species and the production of clonal organs (expressed as total number or per biomass ratio) of three species were higher in NZ than in Germany. When measured around individual plants, the flowering proportion was higher in native German populations of Pilosella officinarum, Hypericum perforatum and Leucanthemum vulgare. Although the study species differed in their invasive status, our findings show that for all six species performance was better in introduced than in native populations. Furthermore, this study emphasizes that multiple measures of plant performance, different spatial scales and differences among species should be taken into account when trying to identify biogeographic differences in the performance of weed species.     

模拟酸雨对入侵植物豚草与伴生种鬼针草竞争关系的影响

植物种间相互作用对物种的生存至关重要。对于入侵植物而言, 更强的种间竞争能力是其成功入侵的重要机制。然而, 环境条件的变化可能会改变种间关系并影响植物入侵的最终结果。该研究主要探索当今严重的环境问题之一——酸雨对中国入侵植物豚草(Ambrosia artemisiifolia)和伴生本地种鬼针草(Bidens bipinnata)种间相互作用的影响, 进一步了解环境扰动对生物入侵的影响。该研究于2021年3月在北京师范大学室外条件下对豚草和鬼针草开展de Wit替代竞争实验, 并施加不同浓度的酸雨模拟溶液(pH = 3、4、5、7), 记录植株在第24、34、45天的株高以及生长季结束时的株高和地上生物量, 通过计算相对邻株效应指数(RNE)并绘制取代系列图表来评估两者的竞争关系。主要结果如下: 单一种植时, 中浓度酸雨(pH = 4)促进了豚草和鬼针草的早期生长, 而高浓度酸雨(pH = 3)显著抑制了两者的早期生长, 但不影响生长季结束时的株高。混合种植时, 高浓度酸雨(pH = 3)显著降低了豚草在第34、45天时的株高, 但在生长季结束时对豚草无影响, 而鬼针草在生长季结束时株高显著降低。鬼针草相对豚草的RNE在任何酸浓度下都不显著。而豚草相对鬼针草的RNE在无酸雨条件下显著, 酸雨处理时在鬼针草比例较低时显著。取代系列实验图表显示, 低浓度酸雨(pH = 5)增加了在低豚草比例下的鬼针草的竞争优势, 而高浓度酸雨增强了豚草的竞争优势。该研究表明酸雨显著影响了豚草和鬼针草的生长及种间竞争关系, 高浓度酸雨增加了入侵植物豚草的竞争优势。     

Projecting potential future shifts in species composition of European urban plant communities

Aim

Urban floras are composed of species of different origin, both native and alien, and with various traits and niches. It is likely that these species will respond to the ongoing climate change in different ways, resulting in future species compositions with no analogues in current European cities. Our goal was to estimate potential shifts in plant species composition in European cities under different scenarios of climate change for the 21st century.

Location

Europe.

Methods

Potential changes in the distribution of 375 species currently growing in 60 large cities in Southern, Central and Western Europe were modelled using generalized linear models and four climate change projections for two future periods (2041–2060 and 2061–2080). These projections were based on two global climate models (CCSM4 and MIROC‐ESM) and two Representative Concentration Pathways (2.6 and 8.5).

Results

Results were similar across all climate projections, suggesting that the composition of urban plant communities will change considerably due to future climate change. However, even under the most severe climate change scenario, native and alien species will respond to climate change similarly. Many currently established species will decline and others, especially annuals currently restricted to Southern Europe, will spread to northern cities. In contrast, perennial herbs, woody plants and most species with temperate continental and oceanic distribution ranges will make up a smaller proportion of future European urban plant communities in comparison with the present communities.

Main conclusions

The projected 21st century climate change will lead to considerable changes in the species composition of urban floras. These changes will affect the structure and functioning of urban plant communities.

     

Differences in leaf thermoregulation and water use strategies between three co‐occurring Atlantic forest tree species

Given anticipated climate changes, it is crucial to understand controls on leaf temperatures including variation between species in diverse ecosystems. In the first study of leaf energy balance in tropical montane forests, we observed current leaf temperature patterns on 3 tree species in the Atlantic forest, Brazil, over a 10‐day period and assessed whether and why patterns may vary among species. We found large leaf‐to‐air temperature differences (maximum 18.3 °C) and high leaf temperatures (over 35 °C) despite much lower air temperatures (maximum 22 °C). Leaf‐to‐air temperature differences were influenced strongly by radiation, whereas leaf temperatures were also influenced by air temperature. Leaf energy balance modelling informed by our measurements showed that observed differences in leaf temperature between 2 species were due to variation in leaf width and stomatal conductance. The results suggest a trade‐off between water use and leaf thermoregulation; Miconia cabussu has more conservative water use compared with Alchornea triplinervia due to lower transpiration under high vapour pressure deficit, with the consequence of higher leaf temperatures under thermal stress conditions. We highlight the importance of leaf functional traits for leaf thermoregulation and also note that the high radiation levels that occur in montane forests may exacerbate the threat from increasing air temperatures.     

BIODIVERSITY RESEARCH: Experimental introduction of the alien plant Hieracium lepidulum reveals no significant impact on montane plant communities in New Zealand

Aim There is debate over whether alien plants necessarily alter the communities they invade or can coexist with native species without discernable impacts. We followed the fate of montane plant communities in response to the experimental sowing of the alien weed Hieracium lepidulum, looking for changes in plant community composition and structure over 6 years. Location Craigieburn Range, New Zealand. Methods We used a replicated randomised block design, with 30 × 30 cm plots (n = 756) subdivided into 5 × 5 cm cells to examine and compare the effects of H. lepidulum at 0.09 m² (plot) and 0.0025 m² (cell) scales. Plots were sown with between 0 and 15,625 H. lepidulum seeds in 2003, forming gradients of invader density and cover. Measurements comprised community richness, evenness and diversity along with H. lepidulum density and cover at both scales. The relationships between the invader and local community attributes were modelled using hierarchical mixed‐effect models. Results Plant communities differed in the extent to which they became invaded, with H. lepidulum cover in the plots ranging from 0% to 52%, with a mean of only 1.89%. Plot species richness increased from 2003 to 2009, with a component of this increase (+0.002 species per year) associated with increasing H. lepidulum density. Other relationships between the plant community and H. lepidulum were generally non‐significant. Main conclusions In these montane plant communities, it appears H. lepidulum coexists with the native community with no measurable negative effects after 6 years on species richness, evenness or diversity, even where density and cover of the invader are highest. We suggest H. lepidulum has persisted preferentially at those sites with abiotic conditions sufficient to support a species‐rich assemblage.     

Interspecific relationships in co‐occurring populations of social parasites and their host ants

Myrmica ant colonies host numerous insect species, including the larvae of Maculinea butterflies and Microdon myrmicae hoverflies. Little is known about the interspecific relationships among these social parasites and their host ants occurring in sympatric populations. We investigated communities of social parasites to assess the strategies allowing them to share the same pool of resources (i.e. Myrmica colonies). The present study was carried out at five sites inhabited by different social parasite communities, each comprising varying proportions of Maculinea teleius, Maculinea nausithous, Maculinea alcon, and Microdon myrmicae. We investigated their spatial distributions, host segregation, the degree of chemical similarity between social parasites and hosts, and temporal overlaps in colony resource exploitation. Spatial segregation among social parasites was found in two populations and it arises from microhabitat preferences and biological interactions. Local conditions can drive selection on one social parasite to use a Myrmica host species that is not exploited by other social parasites. Myrmica scabrinodis and Myrmica rubra nests infested by larvae of two social parasite species were found and the most common co‐occurrence was between Ma. teleius and Mi. myrmicae. The successful coexistence of these two species derives from their exploitation of the host colony resources at different times of the year. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 699–709.