Effects of latitude and soil microbes on the resistance of invasive Solidago canadensis to its co-evolved insect herbivore Corythucha marmorata

纬度和土壤微生物对入侵植物加拿大一枝黄花对共同进化的专性食草昆虫抗性的影响在全球化的背景下,入侵植物再次暴露在来自其原产地的专性食草动物面前的可能性越来越大。然而,入侵植物对曾共同进化的专性食草动物的抗性是否存在纬度趋势以及土壤微生物如何影响入侵植物对这类食草动物的抗性还鲜有研究。我们假设入侵植物加拿大一枝黄花(Solidago canadensis)对其曾共 同进化的来自原产地的食草昆虫菊方翅网蝽(Corythucha marmorata)的抗性随纬度的增加而增加,而加拿 大一枝黄花的局部根际微生物可以促进加其对菊方翅网蝽的抗性。为了验证上述假设,我们首先通过野外调查和温室实验,研究了菊方翅网蝽在中国加拿大一枝黄花种群中的分布丰度,以及对加拿大一枝黄花造成的危害是否存在纬度分布规律。其次,我们通过温室实验,检测了加拿大一枝黄花的本地根际土壤微生物是否能促进加拿大一枝黄花对菊方翅网蝽的抗性。研究结果表明,菊方翅网蝽在加拿大一枝黄花中的分布密度和对加拿大一枝黄花的危害程度与纬度以及加拿大一枝黄花的生长呈现正相关,说明加拿大一枝黄花对菊方翅网蝽的抗性具有纬度趋势。然而,在温室实验中,来自不同纬度的加拿大一枝黄花受到菊方翅网蝽的伤害程度没有显著差异。此外,当根际土壤和加拿大一枝黄花的根状茎同时来自于野外被轻度危害的加拿大一枝黄花或同时来自于野外被重度危害的加拿大一枝黄花时,加拿大一枝黄花的抗性最强。这一结果表明,加拿大一枝黄花的局部根际土壤微生物促进了加拿大一枝黄花对菊方翅网蝽的抗性。     

The dynamics of plant invasions: a case study of three exotic goldenrod species (Solidago L.) in Europe

The patterns of spread of the three exotic species Solidago altissima L., S. gigantea Ait. and S. graminifolia (L.) Salisb. after their introduction to Europe were investigated, based on herbarium specimens and literature records. The spread was analysed by mapping the localities for each decade since 1850. Cumulative numbers of localities as well as numbers of occupied grid squares showed a continuous increase since 1850 for all three species. The slopes, however, were significantly different among the species. Solidago gigantea had the highest colonization rate, followed by S. altissima , and finally S. graminifolia . The latter showed only a slight increase in abundance over time. The increase in diameter of the range in central Europe was logistic for S. altissima and S. gigantea with a rapid increase between 1850 and 1880. The spread of the species in area and time over Europe showed no clear front; new localities at large distances were simultaneously colonized. A large part of the actual range of S. altissima and S. gigantea was already achieved about 1950. It is assumed that the spread of the species followed the hierarchical diffusion model with several independent foci from which the species began to spread. The data suggest that S. altissima and S. gigantea will successfully spread further, leading to an increase in abundance and area, while S. graminifolia seems to spread slowly.     

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.     

Urban landscape and forest vegetation regulate the range expansion of an exotic lace bug Corythucha marmorata (Hemiptera: Tingidae)

Landscapes and vegetation are critical factors in dispersion of exotic insects and expansion of their range. However, few studies have addressed how the surrounding landscape affects the establishment of exotic insects. We assessed the relationship between establishment of an exotic lace bug Corythucha marmorata (Uhler) and the surrounding landscape in the northern edge of the lace bug's expanded range. We found that the lace bugs showed variability in their density among populations. Urban areas had a positive effect, while the natural forest vegetation had a negative effect on lace bug density, with a buffer range of 1–2 km. Moreover, their abundance decreased with distance from the source population. Our results suggest that natural forest landscapes in urban areas may inhibit the range expansion of invasive insects that feed on exotic plants growing in human‐disturbed habitats.     

Role of plant abundance in determining the abundance of herbivorous insects

Summary The proposal (Gaston and Lawton 1988a, b) that small species of insects are more abundant because they have lower per capita resource requirements than large species does not hold for aphids (Dixon 1990a). There are good theoretical grounds, supported by empirical data, for the suggestion that host specific aphids that live on uncommon plants incur great losses in finding their host plants and as a consequence have a lower realized r _m and are rarer than aphids living on common plants. This is also likely to apply to other organisms that are host specific and time-limited dispersers.     

Community structure of insect herbivores on introduced and native Solidago plants in Japan

We compared community composition, density, and species richness of herbivorous insects on the introduced plant Solidago altissima L. (Asteraceae) and the related native species Solidago virgaurea L. in Japan. We found large differences in community composition on the two Solidago species. Five hemipteran sap feeders were found only on S. altissima. Two of them, the aphid Uroleucon nigrotuberculatum Olive (Hemiptera: Aphididae) and the scale insect Parasaissetia nigra Nietner (Hemiptera: Coccidae), were exotic species, accounting for 62% of the total individuals on S. altissima. These exotic sap feeders mostly determined the difference of community composition on the two plant species. In contrast, the herbivore community on S. virgaurea consisted predominately of five native insects: two lepidopteran leaf chewers and three dipteran leaf miners. Overall species richness did not differ between the plants because the increased species richness of sap feeders was offset by the decreased richness of leaf chewers and leaf miners on S. altissima. The overall density of herbivorous insects was higher on S. altissima than on S. virgaurea, because of the high density of the two exotic sap feeding species on S. altissima. We discuss the importance of analyzing community composition in terms of feeding guilds of insect herbivores for understanding how communities of insect herbivores are organized on introduced plants in novel habitats.     

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.     

Habitat expansion of an exotic lace bug, Corythucha marmorata (Uhler) (Hemiptera: Tingidae), on the Kii Peninsula and Shikoku Island in western Japan

Habitat expansion of an exotic lace bug Corythucha marmorata (Uhler) was observed on the Kii Peninsula in 2003–2004, and on Shikoku Island between 2004 and 2007. The bug was first sighted in Nishinomiya City in Hyogo Prefecture in 2000, and then further south in central Kii Peninsula in 2003 before reaching the southern tip of the peninsula by 2004. The bug was first observed in the Tokushima and Ehime Prefectures of Shikoku Island in 2004, after which the range expanded and extended across the entire island by 2007. In this study, six wild Asteraceae species were identified as the host plants to the lace bug.     

浅议我国外来物种入侵问题及其防治对策

随着世界经济的发展和全球经济一体化进程的加快,外来物种入侵已成为影响国家安全的主要因素之一。我国外来入侵物种数量繁多,已成为遭受外来生物危害最严重的国家之一,入侵物种已对我国生态系统、环境和社会经济造成了严重的影响。近年来,各国都积极采取措施,制定外来物种入侵管理政策,加强外来物种入侵立法,确保国家及国际生物安全。然而,我国现有的针对外来物种的相关法律法规存在明显滞后。本文综述了我国外来物种入侵的现状及相关立法,指出外来物种入侵可在不同生态系之间发生,揭示了行政区划与生态系差异造成外来物种入侵管理存在"自然与经济的错位"、国内管理方面存在"生态系统与行政区域的错位"的"双错位现象",分析了我国外来物种的管理现状及今后的发展方向,以期探求我国在国际背景下的外来物种管理对策,提出制定《外来入侵物种法》及相关法律的国家体系、建立跨部门协作科学研究体系以及建设行政管理体系,增强抵御外来物种入侵的能力,实施以防为主的管理策略。     

Establishment of exotic dung beetles in Queensland: the role of habitat specificity

Between 1968 and 1983, 22 species of dung beetles from southern Africa were released into pastoral regions of Queensland in northern Australia. Seven species have become widespread and abundant and the others either occur in isolated patches, are rare or have failed to establish self perpetuating populations. These species all occur in the Hluhluwe district in southern Africa which has a climate similar to that of coastal central and southern Queensland. The numberical abundance and habitat associations of the 22 speices were examined at Hluhluwe to seek an explanation for their performance in Australia. It was concluded that habitat specificity and the adequacy with which habitat was matched in Australia were critical factors. Species which established in Australia were relatively common in cattle dung in Africa where they showed a preference for grassland on clay-loam soils.      

Introduced plants of the invasive Solidago gigantea (Asteraceae) are larger and grow denser than conspecifics in the native range

Introduced plant species that became successful invaders appear often more vigorous and taller than their conspecifics in the native range. Reasons postulated to explain this better performance in the introduced range include more favourable environmental conditions and release from natural enemies and pathogens. According to the Evolution of Increased Competitive Ability hypothesis (EICA hypothesis) there is a trade‐off between investment into defence against herbivores and pathogens, and investment into a stronger competitive ability. In this study, we conducted field surveys to investigate whether populations of the invasive perennial Solidago gigantea Ait (Asteraceae) differ with respect to growth and size in the native and introduced range, respectively. We assessed size and morphological variation of 46 populations in the native North American range and 45 populations in the introduced European range. Despite considerable variation between populations within continents, there were pronounced differences between continents. The average population size, density and total plant biomass were larger in European than in American populations. Climatic differences and latitude explained only a small proportion of the total variation between the two continents. The results show that introduced plants can be very distinct in their growth form and size from conspecifics in the native range. The apparently better performance of this invasive species in Europe may be the result of changed selection pressures, as implied by the EICA hypothesis.     

Drivers of Solidago species invasion in Central Europe—Case study in the landscape of the Carpathian Mountains and their foreground

AimThe invasion process is a complex, context‐dependent phenomenon; nevertheless, it can be described using the PAB framework. This framework encompasses the joint effect of propagule pressure (P), abiotic characteristics of the environment (A), and biotic characteristics of both the invader and recipient vegetation (B). We analyzed the effectiveness of proxies of PAB factors to explain the spatial pattern of Solidago canadensis and S. gigantea invasion using invasive species distribution models.LocationCarpathian Mountains and their foreground, Central Europe.MethodsThe data on species presence or absence were from an atlas of neophyte distribution based on a 2 × 2 km grid, covering approximately 31,200 km² (7,752 grid cells). Proxies of PAB factors, along with data on historical distribution of invaders, were used as explanatory variables in Boosted Regression Trees models to explain the distribution of invasive Solidago. The areas with potentially lower sampling effort were excluded from analysis based on a target species approach.ResultsProxies of the PAB factors helped to explain the distribution of both S. canadensis and S. gigantea. Distributions of both species were limited climatically because a mountain climate is not conducive to their growth; however, the S. canadensis distribution pattern was correlated with proxies of human pressure, whereas S. gigantea distribution was connected with environmental characteristics. The varied responses of species with regard to distance from their historical distribution sites indicated differences in their invasion drivers.Main conclusionsProxies of PAB are helpful in the choice of explanatory variables as well as the ecological interpretation of species distribution models. The results underline that human activity can cause variation in the invasion of ecologically similar species.     

Geographic variation in the evolution and coevolution of a tritrophic interaction

The geographic mosaic theory of coevolution predicts that geographic variation in species interactions will lead to differing selective pressures on interacting species, producing geographic variation in the traits of interacting species (Thompson 2005). We supported this hypothesis in a study of the geographic variation in the interactions among Eurosta solidaginis and its natural enemies. Eurosta solidaginis is a fly (Diptera: Tephritidae) that induces galls on subspecies of tall goldenrod, Solidago altissima altissima and S. a. gilvocanescens. We measured selection on E. solidaginis gall size and shape in the prairie and forest biomes in Minnesota and North Dakota over an 11-year period. Galls were larger and more spherical in the prairie than in the forest. We supported the hypothesis that the divergence in gall morphology in the two biomes is due to different selection regimes exerted by natural enemies of E. solidaginis. Each natural enemy exerted similar selection on gall diameter in both biomes, but differences in the frequency of natural enemy attack created strong differences in overall selection between the prairie and forest. Bird predation increased with gall diameter, creating selection for smaller-diameter galls. A parasitic wasp, Eurytoma gigantea, and Mordellistena convicta, an inquiline beetle, both caused higher E. solidaginis mortality in smaller galls, exerting selection for increased gall diameter. In the forest there was stabilizing selection on gall diameter due to a combination of bird predation on larvae in large galls, and M. convicta- and E. gigantea-induced mortality on larvae in small galls. In the prairie there was directional selection for larger galls due to M. convicta and E. gigantea mortality on larvae in small galls. Mordellistena convicta-induced mortality was consistently higher in the prairie than in the forest, whereas there was no significant difference in E. gigantea-induced mortality between biomes. Bird predation was nonexistent in the prairie so the selection against large galls found in the forest was absent. We supported the hypothesis that natural enemies of E. solidaginis exerted selection for spherical galls in both biomes. In the prairie M. convicta exerts stabilizing selection to maintain spherical galls. In the forest there was directional selection for more spherical galls. Eurytoma gigantea exerted selection on gall shape in the forest in a complex manner that varied among years. We also supported the hypothesis that E. gigantea is coevolving with E. solidaginis. The parasitoid had significantly longer ovipositors in the prairie than in the forest, indicating the possibility that it has evolved in response to selection to reach larvae in the larger-diameter prairie galls.     

Human land use promotes the abundance and diversity of exotic species on Caribbean islands

Human land use causes major changes in species abundance and composition, yet native and exotic species can exhibit different responses to land use change. Native populations generally decline in human‐impacted habitats while exotic species often benefit. In this study, we assessed the effects of human land use on exotic and native reptile diversity, including functional diversity, which relates to the range of habitat use strategies in biotic communities. We surveyed 114 reptile communities from localities that varied in habitat structure and human impact level on two Caribbean islands, and calculated species richness, overall abundance, and evenness for every plot. Functional diversity indices were calculated using published trait data, which enabled us to detect signs of trait filtering associated with impacted habitats. Our results show that environmental variation among sampling plots was explained by two Principal Component Analysis (PCA) ordination axes related to habitat structure (i.e., forest or nonforest) and human impact level (i.e., addition of man‐made constructions such as roads and buildings). Several diversity indices were significantly correlated with the two PCA axes, but exotic and native species showed opposing responses. Native species reached the highest abundance in forests, while exotic species were absent in this habitat. Human impact was associated with an increase in exotic abundance and species richness, while native species showed no significant associations. Functional diversity was highest in nonforested environments on both islands, and further increased on St. Martin with the establishment of functionally unique exotic species in nonforested habitat. Habitat structure, rather than human impact, proved to be an important agent for environmental filtering of traits, causing divergent functional trait values across forested and nonforested environments. Our results illustrate the importance of considering various elements of land use when studying its impact on species diversity and the establishment and spread of exotic species.     

The incidence of exotic species following clearfelling of Eucalyptus regnans forest in the Central Highlands,Victoria

Abstract Invasion by exotic species following clearfelling of Eucalyptus regnans F. Muell. (Mountain Ash) forest was examined in the Toolangi State Forest in the Central Highlands of Victoria. Coupes ranging in age from < 1- to 10-years-old and the spar-stage forests (1939 bushfire regrowth) adjacent to each of these coupes and a mature, 250-year-old forest were surveyed. The dispersal and establishment of weeds was facilitated by clearfelling. An influx of seeds of exotic species was detected in recently felled coupes but not in the adjacent, unlogged forests. Vehicles and frequently disturbed areas, such as roadside verges, are likely sources of the seeds of exotic species. The soil seed bank of younger coupes had a greater number and percentage of seeds of exotics than the 10-year-old coupes and the spar-stage and mature forests. Exotic species were a minor component (< 1% vegetation cover) in the more recently logged coupes and were not present in 10-year-old coupes and the spar-stage and mature forests. These particular exotic species did not persist in the dense regeneration nor exist in the older forests because the weeds were ruderal species (light-demanding, short-lived and short-statured plants). The degree of influence that these particular exotic species have on the regeneration and survival of native species in E. regnans forests is almost negligible. However, the current management practices may need to be addressed to prevent a more threatening exotic species from establishing in these coupes and forests.