Inferring the origin of populations introduced from a genetically structured native range by approximate Bayesian computation: case study of the invasive ladybird Harmonia axyridis

Correct identification of the source population of an invasive species is a prerequisite for testing hypotheses concerning the factors responsible for biological invasions. The native area of invasive species may be large, poorly known and/or genetically structured. Because the actual source population may not have been sampled, studies based on molecular markers may generate incorrect conclusions about the origin of introduced populations. In this study, we characterized the genetic structure of the invasive ladybird Harmonia axyridis in its native area using various population genetic statistics and methods. We found that native area of H. axyridis most probably consisted of two geographically distinct genetic clusters located in eastern and western Asia. We then performed approximate Bayesian computation (ABC) analyses on controlled simulated microsatellite data sets to evaluate (i) the risk of selecting incorrect introduction scenarios, including admixture between sources, when the populations of the native area are genetically structured and sampling is incomplete and (ii) the ability of ABC analysis to minimize such risks by explicitly including unsampled populations in the scenarios compared. Finally, we performed additional ABC analyses on real microsatellite data sets to retrace the origin of biocontrol and invasive populations of H. axyridis, taking into account the possibility that the structured native area may have been incompletely sampled. We found that the invasive population in eastern North America, which has served as the bridgehead for worldwide invasion by H. axyridis, was probably formed by an admixture between the eastern and western native clusters. This admixture may have facilitated adaptation of the bridgehead population.     

Harmonia axyridis in Great Britain: analysis of the spread and distribution of a non-native coccinellid

Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is native to Asia, and was widely introduced as a biocontrol agent of aphids and coccids in Europe and North America. In Europe H. axyridis is considered to be an invasive alien species. Although not known to have been deliberately introduced to Great Britain, it was first recorded there in 2004, in south-east England. Harmonia axyridis arrived in Great Britain by various means, primarily from mainland Europe, but also from Canada. Extensive national and local media coverage, and a dedicated website (), facilitated public involvement in recording H. axyridis in Great Britain; in excess of 4,000 verified records of the species were received between 2004 and 2006. Through detailed mapping, the objective of our study was to quantify and analyse the spread of H. axyridis in its early stages of invasion in Great Britain. Our data shows that between 2004 and 2006, the species spread north through Great Britain at the rate of 58 km year^-1 and west at the rate of 144.5 km year^-1. In England H. axyridis spread north to Yorkshire and west to Cornwall by the end of 2006, and also reached Wales. Recorded occurrence (of one or more H. axyridis individuals at larval, pupal and/or adult stage) in 10 km squares in Great Britain was: 2004—51; 2005—149; 2006—447. Records of juvenile H. axyridis extend through most of the recorded British range of the species, and we present evidence of bi-voltinism in the population in 2006.     

Specialized predation on plataspid heteropterans in a coccinellid beetle: adaptive behavior and responses of prey attended or not by ants

Two plataspid hemipteran species proliferated on Bridelia micrantha(Euphorbiaceae). Colonies of Libyaspis sp., never attendedby ants, developed on branches, while Caternaultiella rugosa lived at the base of the trunks, mostly in association withCamponotus brutus that attends them in carton shelters. Bothplataspid species are prey of the coccinellid beetle Anisolemniatetrasticta, whose larvae always detected them by contact.When attacked the Libyaspis nymphs cowered, so that the hypertrophiedlateral sides of their tergits made contact with the substrate,but the ladybirds slid their long forelegs under these nymphs,lifted them, and bit them on the ventral face. The Caternaultiellanymphs, which do not have hypertrophied extremities of thetergits, tried to escape at contact with the ladybirds, butwere rarely successful. To capture them, the ladybirds eitheradopted the previous behavior or directly grasped then bitthem. We noted a graded aggressiveness in the ants toward theladybirds according to the situation: no aggressiveness onthe tree branches; stopping the ladybirds that approached theshelters where the ants attended Caternaultiella; and fullattack of ladybirds that tried to capture Caternaultiella nymphssituated outside shelters. The latter behavior can emit analarm pheromone that triggers the dispersion of their congenerswhile attracting attending C. brutus workers. Naive workersare not attracted, so we deduce that this behavior is the resultof a kind of learning.     

Harmonia axyridis in Europe: spread and distribution of a non-native coccinellid

Native to Asia, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is considered an invasive alien ladybird in Europe and North America, where it was widely introduced as a biological control agent of aphids and coccids. In Europe, H. axyridis was sold by various biological control companies from 1995 in France, Belgium and the Netherlands, and was also intentionally released in at least nine other countries. It has spread very rapidly, particularly since 2002, and is now regarded as established in thirteen European countries. The established range extends from Denmark in the north to southern France in the south, and from Czech Republic in the east to Great Britain in the west. In this paper we map the spread and distribution of H. axyridis in Europe, and examine the situation on a country-by-country basis. We report first records of the species in five countries; Spain, Sweden, Denmark, Czech Republic and Italy; and first evidence of H. axyridis establishment in the latter three countries. Despite releases of H. axyridis in Portugal, Spain and Greece, there is little evidence of establishment in southern Europe. It is predicted that the spread and increase within Europe will continue and that H. axyridis will become one of the most widely distributed coccinellids in the continent.     

Invasion history and demographic processes associated with rapid morphological changes in the Red‐whiskered bulbul established on tropical islands

The Red‐whiskered bulbul is a very successful invasive bird species. Morphological differences have been reported among individuals inhabiting the humid and dry coasts of Reunion Island, in a 30‐year‐old population. This suggests a capacity for rapid local adaptation which could explain the general invasive success of this species. However, the origin and invasion history of this population is unknown. It is therefore not possible to establish with certainty the cause of these morphological differences. Here, we investigated the invasion history of populations of Red‐whiskered bulbul established on Reunion Island, Mauritius and Oahu (three geographically similar tropical islands) to assess the link between invasion history and morphological changes in these populations. We first assessed the source(s) of the invasive populations. We then compared the morphology of the individuals between the invasive and native populations and between the dry and humid coasts of invaded islands. Finally, we inferred the invasion history of the invasive populations to investigate the role of neutral processes (e.g. founder effect and drift) on morphology. We found that the invasive populations have a similar origin and that the morphology of the individuals in these populations has diverged in a similar way from the native range, suggesting a convergent adaptation to tropical islands. Like on Reunion, we found differences in morphology between the dry and humid coasts on Mauritius. These morphological differences can be explained by invasion history on Reunion but not on Mauritius. Both neutral evolution and adaptation thus shape the morphology of invasive Red‐whiskered bulbuls.     

Seasonal dynamic variation of pollination network is associated with the number of species in flower in an oceanic island community

植物与传粉者相互作用的传粉网络是一个动态的实体,植物开花物候的季节性变化可以在短时间和长时间内重塑其结构。然而,很少有研究考虑到这种季节性动态变化,特别是海洋岛屿群落的传粉网络。本研究探讨了海洋岛屿群落的植物与传粉者间传粉网络的结构是如何随群落内花资源丰富度的季节性变化而动态变化的。利用春夏秋冬四个季节的植物与传粉者间相互作用的数据,分析了四个季节定性的传粉网络结构的动态变化,研究了中国南海西沙群岛的永兴岛群落传粉网络的季节性动态变化。在这四个季节中,分别收集了连续两个月的植物与传粉者相互作用的数据,并计算了四个网络水平的指标来表征传粉网络的总体结构。采用群落差异性统计分析方法,对群落四个季节的网络结构参数进行比较分析,探讨影响这种动态变化格局的潜在因素。同时计算并比较了植物和传粉功能群在物种水平的网络指标的季节动态变化。研究结果表明,永兴岛群落网络水平的特化性和模块化在四个季节的变化均与植物物种丰富度的变化呈明显相反的变化趋势。开花植物种类的增加可能促进了传粉者之间更激烈的竞争,从而导致生态位重叠的增加,引起传粉网络特化性和模块化的下降,反之亦然。进一步分析表明,传粉网络的季节动态变化的内在驱动力是植物与传粉者间连接的重新组合。因此,传粉者之间生态位重叠的季节性变化导致了植物与传粉者间相互作用的重组,从而推动了该群落内植物与传粉者间相互作用的更替变化。在物种水平上,与其它传粉功能群相比,天蛾类传粉者最特化,而蜜蜂科传粉者的物种作用强度最大。因此,在探索孤立的海洋岛屿生态系统以及其它生态系统中的植物与传粉者间相互作用时,应适当考虑到这些新的发现。     

Invasion by alligator weed,Alternanthera philoxeroides,is associated with decreased species diversity across the latitudinal gradient in China

Aims Invasive species occurrence and their effects on biodiversity may vary along latitudes. We examined the occurrence (species cover) and relative dominance (importance value) of invasive alligator weed, Alternanthera philoxeroides, in its terrestrial habitat in China through a large-scale latitudinal field investigation.Methods We established 59 plots along the latitudinal transect from 21°N to 37°N. We recorded species name, abundance, height and individual species coverage of plants in every quadrat. We then measured α-species diversity variations associated with the A. philoxeroides community across the latitudinal range. We also analyzed the effect of latitude on plant species' distributions in this community by using canonical correspondence analysis (CCA).Important findings We found that species cover and importance value of A. philoxeroides increased in areas <35°N, but decreased at higher latitudes. Lower latitudes supported greater species diversity than higher latitudes. Small-scale invasion of A. philoxeroides was associated with higher species diversity, but community diversity was lower when A. philoxeroides species cover exceeded 36%. Community plant species changed from mesophyte to hygrophyte gradually from low to high latitude. Our research suggests that latitude had significant influences on community diversity which interacted with the biotic resistance of a community and impact of invasion. Consequently, A. philoxeroides may become more invasive and have greater negative impacts on community species diversity in higher latitudes as global climate changes.     

Invasion facilitates hybridization with introgression in the Rattus rattus species complex

Biological invasions result in novel species interactions, which can have significant evolutionary impacts on both native and invading taxa. One evolutionary concern with invasions is hybridization among lineages that were previously isolated, but make secondary contact in their invaded range(s). Black rats, consisting of several morphologically very similar but genetically distinct taxa that collectively have invaded six continents, are arguably the most successful mammalian invaders on the planet. We used mitochondrial cytochrome b sequences, two nuclear gene sequences (Atp5a1 and DHFR) and nine microsatellite loci to examine the distribution of three invasive black rat lineages (Rattus tanezumi, Rattus rattus I and R. rattus IV) in the United States and Asia and to determine the extent of hybridization among these taxa. Our analyses revealed two mitochondrial lineages that have spread to multiple continents, including a previously undiscovered population of R. tanezumi in the south‐eastern United States, whereas the third lineage (R. rattus IV) appears to be confined to Southeast Asia. Analyses of nuclear DNA (both sequences and microsatellites) suggested significant hybridization is occurring among R. tanezumi and R. rattus I in the United States and also suggest hybridization between R. tanezumi and R. rattus IV in Asia, although further sampling of the latter species pair in Asia is required. Furthermore, microsatellite analyses suggest unidirectional introgression from both R. rattus I and R. rattus IV into R. tanezumi. Within the United States, introgression appears to be occurring to such a pronounced extent that we were unable to detect any nuclear genetic signal for R. tanezumi, and a similar pattern was detected in Asia.     

Experimental heatwaves facilitate invasion and alter species interactions and composition in a tropical host-parasitoid community

As mean temperatures increase and heatwaves become more frequent, species are expanding their distributions to colonise new habitats. The resulting novel species interactions will simultaneously shape the temperature-driven reorganization of resident communities. The interactive effects of climate change and climate change-facilitated invasion have rarely been studied in multi-trophic communities, and are likely to differ depending on the nature of the climatic driver (i.e., climate extremes or constant warming). We re-created under laboratory conditions a host-parasitoid community typical of high-elevation rainforest sites in Queensland, Australia, comprising four Drosophila species and two associated parasitoid species. We subjected these communities to an equivalent increase in average temperature in the form of periodic heatwaves or constant warming, in combination with an invasion treatment involving a novel host species from lower-elevation habitats. The two parasitoid species were sensitive to both warming and heatwaves, while the demographic responses of Drosophila species were highly idiosyncratic, reflecting the combined effects of thermal tolerance, parasitism, competition, and facilitation. After multiple generations, our heatwave treatment promoted the establishment of low-elevation species in upland communities. Invasion of the low-elevation species correlated negatively with the abundance of one of the parasitoid species, leading to cascading effects on its hosts and their competitors. Our study, therefore, reveals differing, sometimes contrasting, impacts of extreme temperatures and constant warming on community composition. It also highlights how the scale and direction of climate impacts could be further modified by invading species within a bi-trophic community network.     

Invasion history, habitat preferences and phenology of the invasive ladybird Harmonia axyridis in Belgium

Harmonia axyridis Pallas (Coleoptera: Coccinellidae) is native to Asia, and was widely introduced as a biocontrol agent of aphids and coccids in Europe and North America. In Europe, H. axyridis is considered to be an invasive alien species because of its potential to disrupt native ladybird communities. Since 1999, the Belgian Ladybird Working Group mapped all Belgian Coccinellidae and recorded data on substratum plants and habitat. The first feral H. axyridis population in Belgium was recorded in 2001, but the expansion rate is decreasing because it now colonised the whole country. Recorded occupancy in Belgium showed an average rate of increase of 189% between 2002 and 2006. In Belgium, H. axyridis occurred in a wide range of habitats, including those of high conservation value. However, habitat and land cover analysis showed that H. axyridis is more frequently found in urbanised landscapes than in semi-natural landscapes. A niche overlap analysis based on plant use data showed that the potential to affect native species is higher for generalist, deciduous and coniferous tree ladybird species than for heathland and wetland specialist species. Phenology data showed that H. axyridis is able to reproduce later in the year than native species. Based on recorded distribution, ecology and phenology, we discuss the potential of H. axyridis to disrupt native ladybird assemblages in Belgium.     

Determination the population trends of cereal aphids and associated parasitoids by yellow sticky traps with reference to aphid management on wheat

Population trends of cereal aphids and their associated parasitoids inhabiting wheat plantations were monitored by yellow sticky traps. The identified aphids exhibited one seasonal peak for each and were found to be active during the first half of March. Data revealed that, Rhopalosiphum padi L peak occurred one week later than both of Schizaphis graminum (Rondani) and Rhopalosiphum maidis (Fitch). The numbers of aphid complex was recorded more at 90 than at 30 and 60 cm height. Hymenopterous parasitoids activity is synchronised with aphid species, whereas they appeared in small numbers at early and late wheat season and showed their peak on March, with a positive correlation coefficient with aphid populations. The tested compounds (Karate, Biscaya, Match 5% EC, Tracer 24% SC and Neem Azal T/S) showed 100% reduction in aphid numbers after 24 h post application. However, the general reduction percentages indicated 99.31 > 98.32 > 97.97 > 97.06 > 95.66%, by using the abovementioned compounds, respectively.     

The fungal community associated with the ambrosia beetle Xylosandrus compactus invading the mediterranean maquis in central Italy reveals high biodiversity and suggests environmental acquisitions

In summer 2016 a severe infestation of the alien ambrosia beetle Xylosandrus compactus was recorded from the Mediterranean maquis in the Circeo National Park in Central Italy. Trees and shrubs were infested and displayed wilting and necrosis of terminal branches caused by the combined impact of the insect and associated pathogenic fungi. A preliminary screening carried out on captured adults resulted in the isolation of a discrete number of fungal taxa with different life strategies, ranging from true mutualist (e.g. Ambrosiella xylebori) to plant pathogens (Fusarium spp.). In the present study, high-throughput sequencing was applied to determine the total diversity and functionality of the fungal community associated with X. compactus adults collected in the galleries of three Mediterranean woody hosts, Quercus ilex, Laurus nobilis, and Ceratonia siliqua. The effect of season and host in determining the composition of the associated fungal community was investigated. A total of 206 OTUs composed the fungal community associated with X. compactus. Eighteen OTUs were shared among the three hosts, including A. xylebori and members of the Fusarium solani complex. All but two were previously associated with beetles.Sixty-nine out of 206 OTUs were resolved to species level, identifying 60 different fungal species, 22 of which already reported in the literature as associated with beetles or other insects. Functional guild assigned most of the fungal species to saprotrophs and plant pathogens. Effects of seasonality and host on fungal community assemblage were highlighted suggesting the acquisition by the insect of new fungal taxa during the invasion process. The consequences of enriched fungal community on the risk of the insurgence of novel threatful insect–fungus association are discussed considering direct and indirect effects on the invaded habitat.     

Cultivable fungal community associated with the tropical orchid Dichaea andina

The orchid–fungus relationship has been studied since the discovery that the minute seeds of orchids depend on fungi to support the germination process. With the aim of describing the biodiversity of cultivable endophytic and mycorrhizal fungi from the orchid Dichaea andina, we isolated pure fungal cultures from its roots and identified them by sequencing the internal transcribed spacer. We recorded 22 fungal operational taxonomic units belonging to eight orders of Ascomycota: Eurotiales, Hypocreales, Xylariales, Helotiales, Boliniales, Chaetothyriales, Chaetosphaeriales and Pleosporales. The only Basidiomycota isolated belonged to the genus Ceratobasidium from the order Cantharellales, whose members are known as orchid mycorrhizal fungi. At the genus level, we identified 16 genera, the most common of which were Byssochlamys, Camarops, Trichoderma, Cladophialophora, Fusarium and Xylaria; some of them had been reported previously as orchid endophytes. The relevance of endophytic fungi to their hosts is still unclear, but this widely distributed interaction deserves further investigation.     

Invasion of an intermediate predator: Fish population dynamics in a mathematical model of a trophic chain (As applied to Syamozero lake)

A mathematical model is presented for the dynamics of a spatially heterogeneous predator-prey population system; a prototype is the Syamozero lake fish community. We show that the invasion of an intermediate predator can evoke chaotic oscillations in the population densities. We also show that different dynamic regimes (stationary, nonchaotic oscillatory, and chaotic) can coexist. The “choice” of a particular regime depends on the initial invader density. Analysis of the model solutions shows that invasion of an alien species is successful only in the absence of competition between the juvenile invaders and the native species.     

Order of herbivore arrival on wild cabbage populations influences subsequent arthropod community development

In plant–arthropod associations, the first herbivores to colonise a plant may directly or indirectly affect community assembly on that particular plant. Whether the order of arrival of different arthropod species further modulates community assembly and affects plant fitness remains unclear. Using wild Brassica oleracea plants in the field, we manipulated the order of arrival of early‐season herbivores that belong to different feeding guilds, namely the aphid Brevicoryne brassicae and caterpillars of Plutella xylostella. We investigated the effect of herbivore identity and order of arrival on community assembly on two B. oleracea plant populations during two growth seasons. For this perennial plant, we evaluated whether foliar herbivory also affected herbivore communities on the flowers and if these interactions affected plant seed production. Aphid infestation caused an increase in parasitoid abundance, but caterpillars modulated these effects, depending on the order of herbivore infestation and plant population. In the second growth season, when plants flowered, the order of infestation of leaves with aphids and caterpillars more strongly affected abundance of herbivores feeding on the flowers than those feeding on leaves. Infestation with caterpillars followed by aphids caused an increase in flower‐feeding herbivores compared to the reversed order of infestation in one plant population, whereas the opposite effects were observed for the other plant population. The impact on plant seed set in the first reproductive year was limited. Our work shows that the identity and arrival order of early season herbivores may have long‐term consequences for community composition on individual plants and that these patterns may vary among plant populations. We discuss how these community processes may affect plant fitness and speculate on the implications for evolution of plant defences.     

Egg production by the coccinellid Hippodamia convergens fed on two morphs of the green peach aphid, Myzus persicae

Adults of the lady beetle Hippodamia convergens Guérin-Méneville requiring aphids for egg production, were fed suboptimal amounts of apteriform virginoparous larvae or alatiform gynoparous larvae of the green peach aphid, Myzus persicae (Sulz.), and artificial diet ad lib. The beetles initiated oviposition sooner and deposited significantly more eggs when supplied with the same weight of apterous virginoparous morph. The possible nutritional, hormonal, and ecological implications of the findings are discussed.

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Zusammenfassung Die Eiproduktion des Marienkäfers Hippodamia convergens, welcher hierzu Blattläuse als Nahrung benötigt, wurde bei Fütterung mit zwei verschiedenen Morphen der Blattlaus Myzus persicae untersucht. Der Verzehr derselben Menge flügelloser Virginoparen führte im Vergleich zu geflügelten Gynoparen zu früherer und signifikant erhöhter Eiabgabe. Der hormonale und ernährungsphysiologische Hintergrund und die ökologische Bedeutung dieser Ergebnisse werden diskutiert.

     

五指山热带雨林黑桫椤种群及其所在群落特征

黑桫椤(Alsophila podophylla)属国家二级保护植物.作者在海南岛五指山山地雨林和低地雨林分别设立样地对黑桫椤种群及其所在群落进行了调查,分析了热带雨林黑桫椤种群的年龄结构、空间分布格局、数量分布以及群落结构和物种多样性.结果表明,山地雨林和低地雨林中黑桫椤种群分别为稳定型种群和增长型种群,都是完全集群分布.低地雨林的黑桫椤平均密度是山地雨林的1.7倍.土壤pH值对黑桫椤种群密度有负的影响,全磷含量对其有正的影响.山地雨林林地土壤pH值大而全磷含量低,低地雨林刚好相反.土壤酸碱度和全磷含量的差异是导致两地黑桫椤数量差异的原因之一.pH值对黑桫椤平均密度的影响效应是全磷含量的2.5倍,说明pH值是影响黑桫椤种群密度的主要因素.黑桫椤所在群落垂直结构分为3个乔木亚层、2个灌木亚层及草本层.山地雨林黑桫椤所在群落的Simpson指数和Shannon-Wiener指数分别为0.957和5.305,均匀度为0.811;低地雨林黑桫椤所在群落的Simpson指数和Shannon-Wiener指数分别为0.937和5.484,均匀度为0.804.总之,五指山山地雨林和低地雨林的黑桫椤种群年龄结构、分布格局和数量分布都有一定差异,这些差异是由生境差异造成的.热带雨林黑桫椤所在群落结构复杂,物种多样性丰富.     

转植酸酶基因玉米对步甲群落动态的影响

本研究以陷阱法研究转植酸酶基因玉米及其亲本玉米在整个生长季节对地表步甲类群的影响。每个玉米品系设置6块样地, 两个玉米品系交替排列, 每个样地设置2个采样点, 共设置24个样点, 整个玉米生育期取样8次。本研究共采集步甲标本8 012头, 隶属于11属23种, 其中黄斑青步甲Chlaenius micans个体数量分别占个体总数的87.54%, 为玉米田内的主要常见物种, 蠋步甲Dolichus halensis、 后斑青步甲Chlaenius posticalis和单齿蝼步甲Scarites terricola个体数量分别占除黄斑青步甲之外所有步甲个体总数的34.77%, 31.16%和6.21%, 这些物种组成当地玉米田内的常见物种。步甲物种多样性与常见物种个体数量随季节变化明显, 且转植酸酶基因玉米与其亲本玉米田内步甲物种多样性和常见物种个体数量的季节变化趋势相似度较高。重复测量方差分析（Repeated ANOVA）结果表明, 转植酸酶基因玉米对步甲物种丰富度、 物种多度、 香农威纳多样性指数和均匀度指数均没有显著影响; 转植酸酶基因玉米田内的后斑青步甲个体数量明显增加, 但其他常见物种没有显著变化。基于非度量多维度（NMDS）的群落结构分析表明转植酸酶基因玉米与亲本玉米田内的步甲群落结构非常相似。本文结果表明转植酸酶基因玉米的种植对步甲物种多样性及常见物种没有明显影响。