Assessing potential biological control of the invasive plant, tree-of-heaven, Ailanthus altissima

Tree-of-heaven, Ailanthus altissima, is a deciduous tree indigenous to China and introduced into North America and Europe. It is a serious threat to ecosystems in introduced areas, as the plant is very competitive, and also contains allelopathic chemicals that may inhibit growth of surrounding native plants. In addition, the plant contains secondary chemicals that make it unpalatable to some insects. In this paper we assess potential biological control of this plant by reviewing literature associated with natural enemies of the plant from both its native and introduced regions in the world. Our literature surveys revealed that 46 phytophagous arthropods, 16 fungi, and one potyvirus were reported attacking tree-of-heaven, some apparently causing significant damage in China. Two weevils, Eucryptorrhynchus brandti and E. chinensis, are major pests of the plant in China and are reportedly restricted to tree-of-heaven, showing promise as potential biological control agents. Nymphs and adults of a homopteran insect, Lycorma delicatula and larvae of two lepidopteran species, Samia cynthia and Eligma narcissus, may also cause severe damage, but they are not host specific. Two rust fungi, Aecidium ailanthi J. Y. Zhuan sp. nov. and Coleosporium sp. have been reported on tree-of-heaven in China and are also promising potential candidates for biological control of the plant. Nine insect herbivores and 68 fungi are associated with tree-of-heaven in its introduced range in North America, Europe, and Asia. An oligophagous insect native to North America, the ailanthus webworm, Atteva punctella, may be a potential biocontrol agent for the plant. Among the fungal species, Fusarium osysporum f. sp. perniciosum, caused wilt of tree-of-heaven in North America and may have the potential to control the plant, but its non-target effect should be carefully evaluated. Our review indicates that there is potential for using insects or pathogens to control tree-of-heaven.     

丛枝菌根真菌在内的土壤真菌对入侵植物紫茎泽兰的正反馈：证据来自野外实验

包括紫茎泽兰在内的许多外来植物都能够与新入侵生境的丛枝菌根真菌（ AMF）形成互利共生,因此菌根真菌如何调节外来植物种的入侵是当前亟待研究的问题。测定了紫茎泽兰入侵不同阶段（紫茎泽兰呈零星丛状分布于本地植物群落中[部分入侵生境]及紫茎泽兰单优群落形成期[入侵生境]）的土壤化学性状,而后通过野外试验,采用杀真菌剂处理,研究了包括AMF在内的土壤真菌对紫茎泽兰入侵的反馈作用。紫茎泽兰入侵改变了土壤化学性状。施用杀真菌剂降低了紫茎泽兰叶面积、叶片碳、氮、磷、和δ13 C含量。综合分析发现,在紫茎泽兰与本地植物混生群落中,土壤真菌能够增加紫茎泽兰叶片碳和δ13 C含量,但是不能提高紫茎泽兰的光合作用,表明碳和δ13 C含量的提高,不是光合作用的结果,而是通过其他机制实现的。因此可以得出,在部分入侵生境中,碳从土壤或临近植物经由菌丝网向紫茎泽兰转移。紫茎泽兰入侵不同阶段土壤养分的变化利于紫茎泽兰种群建立,同时利于紫茎泽兰借助真菌（尤其是AMF）从土壤或临近植物转移碳,促进种群扩散,这可能是紫茎泽兰入侵的机制之一。     

Differential population responses of native and alien rodents to an invasive predator,habitat alteration and plant masting

Invasive species and anthropogenic habitat alteration are major drivers of biodiversity loss. When multiple invasive species occupy different trophic levels, removing an invasive predator might cause unexpected outcomes owing to complex interactions among native and non-native prey. Moreover, external factors such as habitat alteration and resource availability can affect such dynamics. We hypothesized that native and non-native prey respond differently to an invasive predator, habitat alteration and bottom-up effects. To test the hypothesis, we used Bayesian state-space modelling to analyse 8-year data on the spatio-temporal patterns of two endemic rat species and the non-native black rat in response to the continual removal of the invasive small Indian mongoose on Amami Island, Japan. Despite low reproductive potentials, the endemic rats recovered better after mongoose removal than did the black rat. The endemic species appeared to be vulnerable to predation by mongooses, whose eradication increased the abundances of the endemic rats, but not of the black rat. Habitat alteration increased the black rat''s carrying capacity, but decreased those of the endemic species. We propose that spatio-temporal monitoring data from eradication programmes will clarify the underlying ecological impacts of land-use change and invasive species, and will be useful for future habitat management.     

Environmentally triggered variability in the genetic variance–covariance of herbivory resistance of an exotic plant Solidago altissima

The variability in the genetic variance–covariance (G‐matrix) in plant resistance and its role in the evolution of invasive plants have been long overlooked. We conducted an additional analysis of the data of a reciprocal transplant experiment with tall goldenrod, Solidago altissima, in multiple garden sites within its native range (USA) and introduced range (Japan). We explored the differences in G‐matrix of resistance to two types of foliar herbivores: (a) a lace bug that is native to the USA and recently introduced to Japan, (b) and other herbivorous insects in response to plant origins and environments. A negative genetic covariance was found between plant resistances to lace bugs and other herbivorous insects, in all combinations of garden locations and plant origins except for US plants planted in US gardens. The G‐matrix of the resistance indices did not differ between US and Japanese plants either in US or Japanese gardens, while it differed between US and Japanese gardens in both US and Japanese plants. Our results suggested that the G‐matrix of the plant resistance may have changed in response to novel environmental differences including herbivore communities and/or other biotic and abiotic factors in the introduced range. This may have revealed a hidden trade‐off between resistances, masked by the environmental factors in the origin range. These results suggest that the stability of the genetic covariance during invasion, and the environmentally triggered variability in the G‐matrices of plant resistance may help to protect the plant against multiple herbivore species without changing its genetic architecture and that this may lead to a rapid adaptation of resistance in exotic plants. Local environments of the plant also have a critical effect on plant resistance and should be considered in order to understand trait evolution in exotic plants.     

Contrasting responses of native and alien plant species to soil properties shed new light on the invasion of dune systems

在陆地生态系统中,沿海沙丘特别容易受到外来植物的入侵。许多研究将沙丘生境的入侵归因于人为因素,但对土壤性质和植物性状在植物 入侵中的作用却知之甚少。本研究考察了沙丘系统土壤特征与外来植物入侵的关系,重点研究了土壤养分、土壤盐分和植物功能特征之间的相互作 用。研究地点是马拉诺和格拉多泻湖(北亚得里亚海)的沙质堰洲岛。根据沙丘系统生态梯度上的主要生境(前沙丘、后沙丘和盐沼),在10 个区域内选择100个地块(4 m × 4 m)。在每个地块中记录所有植物物种的发生和丰度,并收集一个土壤核。对每个土壤样品进行土壤质地、电导率(代表土壤盐 分)、有机碳和氮含量分析,并与本地和外来植物的种类数和覆盖度相关联。分析外来和本地物种主要的生殖功能和营养功能性状在生境中的变化。研究结果表明,土壤性质对外来物种库和本地物种库的影响不同,与植物整体多样性密切相关。在后沙丘(最易入侵)的生境中,高的土壤导电性限制了外来物种的数量,但土壤有机碳含量随着外来植物丰度的增加而增加,这也表明植物入侵与土壤之间存在潜在的反馈过程。只有在后沙丘生境中,随着环境条件的改善和植物竞争的加剧,本地植物和外来植物的功能性状谱有显著的趋同。本研究结果表明,在恶劣条件下,只有本地特有植物才能茁壮成长,而在中等条件下,土壤特性梯度与植物特性协同作用,抑制/促进外来植物丰富度。     

An explicit test for the contribution of environmental maternal effects to rapid clinal differentiation in an invasive plant

Population differentiation of alien invasive plants within their non‐native range has received increasingly more attention. Common gardens are typically used to assess the levels of genotypic differentiation among populations. However, in such experiments, environmental maternal effects can influence phenotypic variation among individuals if seed sources are collected from field populations under variable environmental regimes. In the present study, we investigated the causes of an altitudinal cline in an invasive plant. Seeds were collected from Senecio inaequidens (Asteraceae) populations along an altitudinal gradient in southern France. In addition, seeds from the same populations were generated by intra‐population crossings in a climatic chamber. The two seed lots were grown in a common garden in Central Belgium to identify any evidence of environmentally induced maternal effects and/or an altitudinal cline in a suite of life‐history traits. Results failed to detect any environmental maternal effects. However, an altitudinal cline in plant height and above‐ground biomass was found to be independent of the maternal environment.     

Experimental evidence for a delayed response of the above-ground vegetation and the seed bank to the invasion of an annual exotic plant in deciduous forests

Invasions by alien plants significantly affect native biodiversity and ecosystem functioning. We conducted a 5-year field experiment to investigate potential effects of the annual invasive plant Impatiens glandulifera on both the native above-ground vegetation and the soil seed bank in a deciduous forest in Switzerland. Eight years after the establishment of I. glandulifera, we set up plots in patches invaded by the alien plant, in plots from which the invasive plant had been manually removed and in plots which were not yet colonized by the invasive plant. We examined plant species richness, diversity and plant species composition in the above-ground vegetation and soil seed bank in all plots one year and five years after the initiation of the experiment. The 36 plots (3 plot types × 6 replicates × 2 sites) were equally distributed over two forest sites. Neither the native above-ground vegetation nor the soil seed bank was influenced by the presence of I. glandulifera one year after the start of the field experiment. After five years, however, plant species richness of both the above-ground vegetation and the soil seed bank was reduced by 25% and 30%, respectively, in plots invaded by the alien plant compared to plots from which I. glandulifera had been removed or uninvaded plots. Furthermore, plots invaded by the alien plant had a lower total seedling density (reduction by 60%) and an altered plant species composition in the soil seed bank compared to control plots. Our field experiment indicates that negative effects of the annual invasive plant on the native above-ground vegetation and soil seed bank of deciduous forests become visible with a delay of several years.     

Community‐wide impact of an exotic aphid on introduced tall goldenrod

1. The aphid Uroleucon nigrotuberculatum Olive, which is specialised to the tall goldenrod, Solidago altissima L., in its native range, has become a dominant species on the introduced tall goldenrod in Japan. How this exotic aphid influenced arthropod communities on the introduced tall goldenrod in aphid‐present (spring) and aphid‐absent (autumn) seasons was examined, using an aphid removal experiment. 2. In spring, aphid presence increased ant abundance because aphid honeydew attracted foraging ant workers. A significant negative correlation was found between the numbers of ants and herbivorous insects other than aphids on the aphid‐exposed plants, but no significant correlation was detected on the aphid‐free plants. Thus, the aphid presence was likely to decrease the abundance of co‐occurring herbivorous insects through removal behaviour of the aphid‐tending ants. There were no significant differences in plant traits between the aphid‐exposed and aphid‐free plants. 3. In autumn, the numbers of lateral shoots and leaves, and the leaf nitrogen content were increased in response to the aphid infestation in spring. Because of the improvement of plant traits by aphid feeding, the abundance of leaf chewers increased on aphid‐exposed plants. In contrast, the abundance of sap feeders decreased on the aphid‐exposed plants. In particular, the dominant scale insect among sap feeders, Parasaissetia nigra Nietner, decreased, followed by a decrease in the abundance of ants attending P. nigra. Thus, aphid feeding may have attenuated the negative impacts of the tending ants on leaf chewers. 4. Aphid presence did not change herbivore species richness but changed the relative density of dominant herbivores, resulting in community‐wide effects on co‐occurring herbivores through ant‐mediated indirect effects, and on temporally separated herbivores through plant‐ and ant‐mediated indirect effects. The aphid also altered predator community composition by increasing and decreasing the relative abundance of aphid‐tending ants in the spring and autumn, respectively.     

PhenoSpace: A Shiny application to visualize trait data in the phenotypic space of the global spectrum of plant form and function

1. A recent analysis of variation in six major traits conducted on a large worldwide sample of vascular plant species showed that three‐quarters of trait variation was captured by a two‐dimensional global spectrum of plant form and function (“global spectrum” hereafter). We developed the PhenoSpace application, whose aim is to visualize and export the position of any individual/population/species in the phenotypic space of the global spectrum.
2. PhenoSpace is a Shiny application that helps users to manipulate and visualize data pertaining to the global spectrum of plant form and function. It is freely accessible at the following URL: https://shiny.cefe.cnrs.fr/PhenoSpace/.
3. PhenoSpace has three main functionalities. First, it allows users to visualize the phenotypic space of the global spectrum using different combinations of traits and growth forms. Second, trait data from any new user‐defined dataset can be projected onto the phenotypic space of the global spectrum, provided that at least two of the six traits are available. Finally, figures produced and loadings of the imported data on the PCA axes can be downloaded, allowing users to conduct further analyses.
4. PhenoSpace fulfills the practical goal of positioning plants in the phenotypic space of the global spectrum, making it possible to compare trait variation at any level of organization against the worldwide background. This serves a major aim of comparative plant ecology, which is to put specific sets of individuals, populations or species into a broader context, facilitating comparison and synthesis of results across different continents and environments using relevant indicators of plant design and function.

     

加拿大一枝黄花在安徽合肥的入侵生态因子分析

【目的】研究加拿大一枝黄花入侵区域的入侵格局,分析其入侵性与当地生态环境之间的关系。【方法】通过随机布设样地方法量化了该植物入侵区域的海拔、坡度、植物群落等22个生境因子,并进行主成分分析。【结果】(1)加拿大一枝黄花生境的22个生态因子所含的生态信息可用6个综合因子表征:土壤肥力因子、土壤理化因子、植被因子、复合因子、干扰因子和光源因子;(2)加拿大一枝黄花的发生频率与速效钾、样地中植物总株数、杂草盖度、有机质、植物种数、海拔、坡度、乔木高度、乔木大小、土壤含水率等生态因子呈正相关,但是与水源距、土壤有效磷、土壤全磷、土壤pH呈负相关。【结论】加拿大一枝黄花入侵可能与当地自然环境中的某些生态因子密切相关,生态因子可能对其入侵成功产生抑制或促进作用。本研究有助于明确加拿大一枝黄花入侵的生态因子,为该入侵植物的有效防治提供一定的理论依据。     

Variation in host plant has no effect on the performance and fitness‐related traits of the specialist herbivore Pareuchaetes insulata

The effect of host plant dissimilarity on insect preference and performance was tested using two morphological forms of Chromolaena odorata (L.) King & Robinson (Asteraceae) (one from Florida, USA, another from South Africa), and a specialist herbivore, Pareuchaetes insulata (Walker) (Lepidoptera: Erebidae: Arctiinae) from Florida, USA, that was introduced as a biological control agent in South Africa. Although this insect did establish at one site, of some 21 sites at which over 800 000 individuals were released, its population level in the field has remained low after an initial outbreak in 2006. To explain the poor performance of P. insulata, we hypothesised that P. insulata larvae prefer Floridian C. odorata to the southern African C. odorata, which is morphologically and probably genetically distinct, and that larvae reared on Floridian C. odorata should have higher fitness and performance. We tested this by comparing insect performance metrics on each of the two plant forms in laboratory experiments. Apart from pupal mass, which was significantly greater on southern African C. odorata, P. insulata performance metrics were similar on both plant forms; there were no significant differences in total leaf area consumed, egg and larval development, immature survival rates, feeding index, host suitability index, growth index, and fecundity between the Floridian and southern African C. odorata plants. In sum, we could not demonstrate that differences in plant forms in C. odorata are responsible for the poor performance of P. insulata in South Africa.     

Integrating species distribution models and interacting particle systems to predict the spread of an invasive alien plant

Aim We demonstrate how to integrate two widely used tools for modelling the spread of invasive plants, and compare the performance of the combined model with that of its individual components using the recent range dynamics of the invasive annual weed Ambrosia artemisiifolia L. Location Austria. Methods Species distribution models, which deliver habitat‐based information on potential distributions, and interacting particle systems, which simulate spatio‐temporal range dynamics as dependent on neighbourhood configurations, were combined into a common framework. We then used the combined model to simulate the invasion of A. artemisiifolia in Austria between 1990 and 2005. For comparison, simulations were also performed with models that accounted only for habitat suitability or neighbourhood configurations. The fit of the three models to the data was assessed by likelihood ratio tests, and simulated invasion patterns were evaluated against observed ones in terms of predictive discrimination ability (area under the receiver operating characteristic curve, AUC) and spatial autocorrelation (Moran’s I). Results The combined model fitted the data significantly better than the single‐component alternatives. Simulations relying solely on parameterized spread kernels performed worst in terms of both AUC and spatial pattern formation. Simulations based only on habitat information correctly predicted infestation of susceptible areas but reproduced the autocorrelated patterns of A. artemisiifolia expansion less adequately than did the integrated model. Main conclusions Our integrated modelling approach offers a flexible tool for forecasts of spatio‐temporal invasion patterns from landscape to regional scales. As a further advantage, scenarios of environmental change can be incorporated consistently by appropriately updating habitat suitability layers. Given the susceptibility of many alien plants, including A. artemisiifolia, to both land use and climate changes, taking such scenarios into account will increasingly become relevant for the design of proactive management strategies.     

放牧强度对高寒草甸土壤理化性状和植物功能群的影响

为明晰放牧强度对植物功能群落的物种分布特征和土壤理化性状的影响。在青藏高原东北缘高寒草甸设置6块放牧强度样地开展试验。采用多元排序和方差分解等方法,分析放牧强度作用下植物功能群落物种分布与土壤物理结构和化学养分因子的定量关系。结果表明:(1)随放牧强度增加,禾本科、莎草科和多年生杂类草植物群落的物种丰富度、盖度和实际重要值均显著降低,而一年生杂类草无显著变化。(2)对于土壤化学养分,不同放牧强度区,土壤速效钾、全氮和有机质含量均随土层深度的增加而降低,土壤速效磷在不同放牧强度区变化规律不一致。随放牧强度增加,土壤速效钾和全氮含量增加,而土壤速效磷和有机质含量无明显变化。(3)对于土壤物理性状,不同放牧强度区,随土层深度增加,土壤紧实度和容重均增加,而土壤含水量和通气孔隙度均降低。随放牧强度增加,0—30 cm各土层深度的土壤紧实度和容重均增加,土壤含水量和通气孔隙度均降低。(4)以功能群为基本单元对植物群落进行方差分解得出,土壤物理性状可单独解释群落功能群分布总方差的58.10%。(5)基于物种尺度对不同功能群落与环境因子进行定量研究表明,土壤紧实度是决定每种功能群物种分布格局的最主要因子。禾本科、莎草科和多年生草本植物均受土壤物理性状的显著影响,贡献率分别为26.3%、31.0%和16.5%。而一年生草本植物不受土壤物理和化学性状的显著影响。综上所述,放牧强度对土壤化学性状的影响具有不确定性,而对土壤物理性状和群落特征的影响具有确定性。土壤物理结构性状主导了高寒草甸植物功能群落的物种分布格局。     

The potential for birds to disperse the seeds of Acacia cyclops,an invasive alien plant in South Africa

Rooikrans Acacia cyclops is an aggressive invasive tree that threatens natural resources in South Africa. The seeds of A. cyclops have a prominent aril which attracts birds that ingest the seeds and disperse them endozoochorously. Two biological control agents, a Seed Weevil Melanterius servulus and a Flower‐galling Midge Dasineura dielsi, were released on A. cyclops in 1991 and 2002, respectively. Together these agents have substantially reduced seed production and generally far lower numbers of seeds are now available to birds. A consequence of this transition from historically bounteous quantities of seeds to scanty seed availability is that birds may no longer associate with the trees and seed dispersal may be disproportionately reduced. To assess whether this has happened, seed attrition was measured by comparing the amount of seeds that disappeared from two groups of branches, one available to birds and the other enclosed in bird netting. Other types of granivores (mainly field mice) were excluded from both groups of branches with a plastic funnel placed around the stems. Mature seeds were also harvested and fed to caged bird species to determine gut retention times and germination rates of ingested seeds. Attrition rates of seeds showed that birds continue to remove seeds but that only a proportion of the crop is taken. Only two frugivorous species (Knysna Turaco Tauraco corythaix and Red‐winged Starling Onychognathus morio) and two granivorous species (Red‐eyed Dove Streptopelia semitorquata and Laughing Dove Streptopelia sengalensis) ingested A. cyclops seeds during feeding trials. Ingestion by birds enhanced seed germination except for those ingested by Laughing Doves. There were no apparent effects of length of gut passage time and avian body size on seed germination rates. Despite the diminished seed resource due to biological control agents, birds continue to disperse A. cyclops seeds.     

Factors that shape the elevational patterns of plant diversity in the Yatsugatake Mountains,Japan

Elevation is involved in determining plant diversity in montane ecosystems. This study examined whether the distribution of plants in the Yatsugatake Mountains, central Japan, substantiated hypotheses associated with an elevational diversity gradient. Species richness of trees, shrubs, herbs, ferns, and bryophytes was investigated in study plots established at 200‐m elevational intervals from 1,800 to 2,800 m. The changes in plant diversity (alpha and beta diversities, plant functional types, and elevational ranges) with elevation were analyzed in relation to climatic factors and elevational diversity gradient hypotheses, that is, mass effect, mid‐domain effect, and Rapoport''s elevational rule. In addition, the elevational patterns of dominance of plant functional types were also analyzed. A comparison of alpha and beta diversities revealed that different plant groups responded variably to elevation; the alpha diversity of trees and ferns decreased, that of herbs increased, whereas the alpha diversity of shrubs and bryophytes showed a U‐shaped relationship and a hump‐shaped pattern. The beta diversity of shrubs, herbs, and bryophytes increased above the subalpine–alpine ecotone. In accordance with these changes, the dominance of evergreen shrubs and graminoids increased above this ecotone, whereas that of evergreen trees and liverworts decreased. None of the plant groups showed a wide elevational range at higher elevations. These elevational patterns of plant groups were explained by climatic factors, and not by elevational diversity gradient hypotheses. Of note, the changes in the dominance of plant groups with elevation can be attributed to plant–plant interactions via competition for light and the changes in physical habitat. These interactions could alter the elevational diversity gradient shaped by climatic factors.