Intra- and interspecific competition between biotypes B and Q of Bemisia tabaci (Hemiptera: Aleyrodidae) from Spain

Biotypes B and Q of Bemisia tabaci (Gennadius), collected from the islands of Tenerife and Majorca respectively, were exposed to competition conditions on tomato cv. Marmande in the laboratory. Both biotypes were established in single and mixed cultures at different densities. Increased mortality of biotype Q females and immature instars was observed together with a lower rate of fecundity and progeny size compared to biotype B, when reared in single or mixed cultures. The female:male sex ratio of F1 individuals of biotype Q was higher in single than in mixed cultures. However, the sex ratio of F1 individuals of biotype B was the same in single and mixed cultures, suggesting reproductive interference. Whitefly density did not affect interspecific interactions. It had a moderate effect on developmental rate of both biotypes, and on mortality of immature instars and progeny size of biotype B only. The results indicate that under laboratory conditions the studied biotype B should displace biotype Q.     

Crossing experiments and behavioral observations reveal reproductive incompatibility among three putative species of the whitefly Bemisia tabaci

Abstract The whitefly Bemisia tabaci has a global distribution and extensive genetic diversity. Recent phylogenetic analyses as well as crossing experiments suggest that B. tabaci is a complex composed of > 20 cryptic species, but more crossing studies are required to examine the reproductive compatibility among the putative species and thus further clarify the systematics of this species complex. We conducted crossing experiments and behavioral observations to investigate the reproductive compatibility between the Mediterranean, Asia II 3, and Asia II 1 putative species of B. tabaci collected from Zhejiang, China. Female progeny were never produced in inter-species crosses, demonstrating a lack of egg fertilization; while 55%–75% females were produced in all the intra-species treatments. Continuous behavioral observations showed that frequent courtship events occurred in both intra-species treatments and inter-putative species crosses. However, copulation events occurred only in the three intra-species treatments with one exception: that one copulation event occurred between Asia II 3 and Mediterranean in the crosses where two cohorts of females and males of different putative species were enclosed together in a small arena but were not allowed access to their intra-specific mates for a long period of time. These data demonstrated complete reproductive isolation between the Mediterranean, Asia II 3, and Asia II 1 putative species, and further showed that the isolation is due to lack of copulation. Demonstration of reproductive isolation between the Mediterranean and two indigenous putative species from China provides further evidence for the existence of cryptic species within the B. tabaci complex.     

Genetic mutations associated with chemical resistance in the cytochrome P450 genes of invasive and native Bemisia tabaci (Hemiptera: Aleyrodidae) populations in China

Abstract  Bemisia tabaci (Gennadius) is a species complex, and its two most damaging biotypes B and Q are globally distributed pests. Despite increasing biological and economic impacts, little is known about the evolutionary mechanisms that favor their competition with native populations. Here, we investigated the genetic mutations in the P450 gene of the invasive B, Q biotypes and the native Cv population. Four mutations associated with chemical resistance, Pro-Leu, Ala-Ser, Ser-Phe and Trp-Leu, were found in the cytochrome P450 CYP6C and CYP9F genes of the B and Q biotypes. Bioassay results also revealed that both the B and Q biotypes have about 12–47 times more resistance to acephate, beta-cypermethrin, methomyl, and 5–7 times more resistance to imidacloprid insecticide than Cv population. Our results provide a molecular approach for better understanding and monitoring the pesticide resistances of invasive and native B. tabaci populations in China.     

Introgression of resistance genes between populations: a model study of insecticide resistance in Bemisia tabaci

Introgression is a key process in conservation biology, genetic modification of (crop) species and in the evolutionary ecology of many species. Here we consider the case of introgression of insecticide resistance in the whitefly, Bemisia tabaci. B. tabaci is a species complex consisting of a range of biotypes, known to have a high degree of inter-biotype reproductive isolation. In areas where insecticide resistant and susceptible biotypes of B. tabaci coexist, introgression of the resistance gene will have considerable consequences for whitefly control. Using a stochastic branching process model we calculate the relative importance of life-history traits in determining the probability of introgression given that a hybridization event has occurred. We show that a fitness cost expressed through the average number of eggs laid, has the largest effect on the introgression probability as compared to fitness costs expressed through other life-history parameters. These results change when we consider a reproductive isolation mechanism, for which we show that the fitness cost expressed through the male survival and mating probability have the largest effect on the probability of introgression.     

浙江B型与一非B型(China-ZHJ-1)烟粉虱形态学和生物学特性的比较研究

本文对新入侵我国的B型与浙江一非B型（China-ZHJ-1种群）烟粉虱之间的生殖亲和性进行了测定,并对两个生物型在形态特征、发育历期、诱导西葫芦银叶反应及寄主植物适应性几方面进行了比较研究。结果表明,当将B型与非B型ZHJ-1种群进行杂交组合时,两生物型之间虽然有求偶行为发生,但从不发生交配,后代全部为孤雌生殖产下的雄虫;而B型、非B型各自内部羽化后0～72 h的平均交配次数分别为4.4次和1.0次, 所产后代的雌雄性比分别为2.8∶1、1.0∶1,前者性比显著高于后者。两生物型之间伪蛹形态存在明显差异,非B型ZHJ-1种群的前、后蜡缘饰宽度分别为B型的2.5～2.7倍、1.8～2.1倍。在棉花上,除B型1龄若虫历期较非B型的略长外,两生物型之间其他各虫态的发育历期均无显著差异;从2龄若虫起,非B型各虫态的体长明显长于B型,并且同一生物型内,4龄雌性若虫个体明显大于同龄雄性若虫。B型在多种寄主植物上的存活力比非B型的显著要强。B型与非B型ZHJ-1种群生物学特性的比较,为对我国烟粉虱这两个生物型进行深入的比较研究、探讨B型烟粉虱的入侵生物学提供了基础。     

不同作物田烟粉虱发生的时空动态

2008～2009年连续2年系统调查了番茄、茄子、棉花、大豆、玉米等寄主植物上烟粉虱种群发生的时间与空间动态。结果表明：不同寄主植物上的烟粉虱成虫及其伪蛹数量有显著性差异,其密度大小依次为：茄子>棉花>番茄>大豆>玉米。其中,在玉米上除了发现极少量的成虫逗留外,没有发现烟粉虱的卵及若虫。在发生的时间序列上,烟粉虱成虫及伪蛹的数量呈现为先逐渐上升后又下降的变化过程,发生高峰期集中在8月5日到8月31日,9月初以后烟粉虱数量慢慢减少。在空间分布上,表现为烟粉虱成虫喜食寄主的上部叶片。统计分析显示,寄主对烟粉虱成虫和伪蛹的数量的影响极显著,而年份对其数量的影响没有显著差异。由此得出的烟粉虱发生和达到高峰的时间,可为烟粉虱预测预报和区域性综合治理提供重要理论依据。     

微卫星位点BEM06与BEM23鉴别烟粉虱B型与Q型的有效性

烟粉虱Bemisia tabaci (Gennadius) B型与Q型是烟粉虱复合种中入侵性较强、分布较广的2种生物型, 当前在许多地区混合发生。这2种生物型的快速鉴别对其种群动态调查及入侵生态学研究具有重要价值。为了验证微卫星位点BEM06与BEM23鉴别B型与Q型烟粉虱的有效性, 本研究分析了这2个微卫星位点的等位基因在国内外17个B型、4个Q型、3个非B/Q型烟粉虱种群的分布特点。结果表明: 这2个微卫星位点的联合使用可鉴别B型与Q型烟粉虱, 但是无法有效地将B型、Q型与一些非B/Q型烟粉虱某些个体区分开来。结果提示: 利用微卫星位点BEM06与BEM23鉴别B型与Q型烟粉虱具有一定的局限性, 尤其是田间烟粉虱存在其他生物型时需要慎重使用这种鉴别方法。     

烟粉虱B型与Q型种群遗传多样性的AFLP分析

烟粉虱Bemisia tabaci(Gennadius)是由多种生物型或物种组成的复合种。Q型烟粉虱在我国部分地区正在取代B型烟粉虱成为优势生物型。烟粉虱Q型与B型遗传多样性比较研究为解析这2种生物型入侵的遗传学基础具有重要的意义。本文利用AFLP技术研究了Q型(包括Q1型、Q2型)、B型烟粉虱种群的遗传多样性。结果表明:Q型烟粉虱遗传多样性高于B型烟粉虱;Q1型烟粉虱各项遗传多样性指数均接近于Q2型。最后探讨了AFLP与SSR分子标记的各自特点。     

mtCOI PCR-RFLP技术鉴别中国入侵型和土著型烟粉虱种群的有效性分析

烟粉虱Bemisia tabaci (Gennadius)是世界范围内最重要的入侵生物之一,准确地鉴别烟粉虱入侵生物型和土著生物型具有十分重要的现实意义。mtCOI PCR-RFLP技术具有快速、高效的特点,是当前应用最广泛的烟粉虱生物型鉴定技术,但是,不同限制性内切酶为基础的mtCOI PCR-RFLP技术在鉴定我国烟粉虱种群中的有效性仍不明了。本文比较了已报导的5种mtCOI PCR-RFLP技术在鉴别中国烟粉虱种群入侵生物型和土著生物型(B型、Q型,ZHJ1型、ZHJ2型、ZHJ3型)的有效性。结果表明,内切酶AluI不能区分B型和ZHJ2型烟粉虱;内切酶TaqI不能准确区分ZHJ3型和Q型烟粉虱个体;而内切酶VspI不仅不能准确区分ZHJ1型和Q型烟粉虱个体,也不能准确区分B型和ZHJ2型烟粉虱;内切酶MseI和Tru9I则不能有效鉴别上述5种烟粉虱生物型,因此不适宜推广使用。     

Evidence from molecular markers and population genetic analyses suggests recent invasions of the Western North Pacific region by biotypes B and Q of Bemisia tabaci (Gennadius)

Invasive events by Bemisia tabaci (Gennadius) biotypes in various parts of the world are of continuing interest. The most famous is biotype B that has caused great economic losses globally. In addition, biotype Q has also recently been reported to be a new invasive pest. These two biotypes have been monitored for some time in the Western North Pacific region, but the invasive events and population genetic structures of these two biotypes are still not clear in this region. In this study, the mitochondrial cytochrome oxidase I (COI) gene was used to reconstruct a phylogenetic tree for identifying biotypes B and Q and to study the relationships between invasive events and ornamental plants. Population genetic analyses of mtCOI sequences were also used to study the genetic relationships within and between populations. A combination of a phylogenetic tree and haplotype analysis suggested the recent invasion of biotype Q in this region is related to the international ornamental trade from the Mediterranean region. Low levels of haplotype diversity and nucleotide diversity indicate that the presence of biotypes B and Q in the Western North Pacific region are caused by multiple invasions. Hierarchical analysis of molecular variance supports the hypothesis of multiple invasions. In addition, high sequence identities and low genetic distances within and between populations of the two biotypes revealed that these invasive events occurred recently. The low levels of genetic differentiation revealed by pairwise F (ST) values between populations also suggests the invasions were recent. Therefore, results of this study suggested that biotypes B and Q entered this region through multiple recent invasions. A quarantine of agricultural crops may be necessary to prevent further invasions.     

寄主型和迁飞型棉蚜的交配行为

棉蚜Aphis gossypii Glover种群存在寄主利用和迁飞能力上的明显分化,有棉花型和瓜型、迁飞型和滞留型之分。但是,棉花型与瓜型之间,以及迁飞型与滞留型之间是否能发生交配行为,尚无研究报道。本文在低温和短光照条件下分别诱导棉花型、瓜型、迁飞型和滞留型棉蚜的性蚜,并进行性蚜间的交配行为观察。结果表明,迁飞型和滞留型性蚜间可以发生交配行为,杂交时发生交配行为的个体比率与自交时无显著差异,但是杂交时雄蚜寻找配偶所花的时间要显著长于自交时,并且迁飞型雌蚜(M♀)与滞留型雄蚜(S♂)杂交时的交配持续时间也显著长于自交时。迁飞型和滞留型棉蚜同型交配容易完成,棉蚜的迁飞型和滞留型已在交配行为上产生了一定的分化。棉花型和瓜型棉蚜的性蚜间可以发生交配行为,并且两品系在正交和反交时雄蚜的寻偶时间与交配持续时间均无显著差异。     

辽宁省冬季温室内越冬粉虱伪蛹的种类鉴定及烟粉虱体内番茄黄化曲叶病毒的检测

为了揭示辽宁省冬季温室内越冬粉虱伪蛹的种类及烟粉虱Bemisia tabaci （Gennadius）携带番茄黄化曲叶病毒（tomato yellow leaf curl virus, TYLCV）情况, 于2012年1月份在辽宁省不同县市区的温室作物上采集了17份粉虱伪蛹样品（每样品含30头粉虱伪蛹） , 镜检鉴别粉虱种类并利用mtCOI基因对烟粉虱生物型进行了鉴定; 检测了烟粉虱携带TYLCV情况并对其PCR扩增产物进行了测序分析。结 果表明： 辽宁省冬季温室内存在越冬温室白粉虱Trialeurodes vaporariorum （Westwood）与烟粉虱。17份粉虱样品中, 11份样品为烟 粉虱样品, 6份样品为温室白粉虱和烟粉虱混合样品。混合样品中, 温室白粉虱仅在锦州凌海（LH）样品中占优势。17份烟粉虱样品（包 括混合样品）中, 仅有4份样品为B型与Q型混合样品, 其他13份样品烟粉虱生物型均为Q型。17份烟粉虱样品中有3份Q型烟粉虱样品检测 到TYLCV, 系统树分析进一步证实该病毒是TYLCV。调查结果为辽宁省粉虱与TYLCV的早期测报和防控提供了科学依据。     

华东、华南和西南地区烟粉虱的种群及寄主分化

烟粉虱Bemisia tabaci是一个复合种,它具有的生物型分化、较强的传播病毒的能力和抗药性、较快的繁殖速率等特征使其成为我国农业生产中重要害虫之一。本研究利用细胞色素线粒体氧化酶Ⅰ基因,对采集自江苏、广东和云南三省的烟粉虱样本进行了生物型鉴定,并对烟粉虱生物型与寄主植物之间的关联性开展了调查。结果表明,在广东和云南省,都存在未鉴定的土著种群与入侵的B型、Q型共存的现象;同时,在本研究中广东省尚未采集到Q型烟粉虱,而在江苏采集到的粉虱样本全部为入侵型。研究结果还表明,相对于入侵种而言,土著种群显示出更强的寄主植物趋同性;丰富的寄主植物以及本身具有的多食性特性有助于B型、Q型等生物型在世界各地的广泛入侵。     

Biotype status and distribution of Bemisia tabaci (Hemiptera: Aleyrodidae) in Shandong province of China based on mitochondrial DNA markers

Bemisia tabaci has caused significant crop losses in China during the last decade. Recent research has shown that two potentially invasive variants, biotypes B and Q, have been found in several regions of China. Our objective was to determine the biotype status and the distribution of B. tabaci in Shandong province, an important agricultural region of China. Based on mitochondrial DNA markers, both biotypes B and Q were detected, with B being the predominant biotype. The results indicate that the more recently introduced biotype Q has not only been located in China but also has established and spread in some regions.     

烟粉虱生物型的监测及其遗传结构研究

烟粉虱Bemisia tabaci(Gennadius)是一种重要的农业害虫并包括许多生物型,其中B型和Q型是入侵性较强的2种生物型。文章着重介绍近年来在烟粉虱生物型的监测及其遗传结构方面的研究进展。B型烟粉虱和Q型烟粉虱这2个生物型均已入侵我国,其中多数地区烟粉虱是B型烟粉虱,局部地区有Q型烟粉虱并呈现不断蔓延趋势。微卫星(SSR)分子标记分析结果表明我国B型烟粉虱的入侵来源具有多元化,而云南地区Q型烟粉虱来源比较单一。化学农药的使用能够影响室内种群的遗传结构,降低种群的遗传多样性。基于RAPD、ISSR分子标记的分析结果表明,Q型烟粉虱种群各项遗传多样性指数均比B型烟粉虱的高。今后加强烟粉虱入侵生物型的遗传结构及其种群动态关系等方面的研究,对于揭示烟粉虱的灾变机制及其控制具有重要的意义。     

Invasive mechanism and management strategy of Bemisia tabaci (Gennadius) biotype B: Progress report of 973 Program on invasive alien species in China

Bemisia tabaci (Gennadius) biotype B, called a “superbug”, is one of the most harmful biotypes of this species complex worldwide. In this report, the invasive mechanism and management of B. tabaci bio-type B, based on our 5-year studies, are presented. Six B. tabaci biotypes, B, Q, ZHJ1, ZHJ2, ZHJ3 and FJ1, have been identified in China. Biotype B dominates the other biotypes in many regions of the country. Genetic diversity in biotype B might be induced by host plant, geographical conditions, and/or insecticidal application. The activities of CarE (carboxylesterase) and GSTs (glutathione-S-transferase) in biotype B reared on cucumber and squash were greater than on other host plants, which might have increased its resistance to insecticides. The higher activities of detoxification enzymes in biotype B might be induced by the secondary metabolites in host plants. Higher adaptive ability of biotype B adults to adverse conditions might be linked to the expression of heat shock protein genes. The in-digenous B. tabaci biotypes were displaced by the biotype B within 225 d. The asymmetric mating in-teractions and mutualism between biotype B and begomoviruses via its host plants speed up wide-spread invasion and displacement of other biotypes. B. tabaci biotype B displaced Trialeurodes vapo-rariorum (Westwood) after 4－7 generations under glasshouse conditions. Greater adaptive ability of the biotype B to adverse conditions and its rapid population increase might be the reasons of its suc-cessful displacement of T. vaporariorum. Greater ability of the biotype B to switch to different host plants may enrich its host plants, which might enable it to better compete with T. vaporariorum. Native predatory natural enemies possess greater ability to suppress B. tabaci under field conditions. The kairomones in the 3rd and 4th instars of biotype B may provide an important stimulus in host searching and location by its parasitoids. The present results provide useful information in explaining the mechanisms of genetic diversity, evolution and molecular eco-adaptation of biotype B. Furthermore, it provides a base for sustainable management of B. tabaci using biological and ecological measures.     

Multiple origins of pyrethroid resistance in sympatric biotypes of Bemisia tabaci (Hemiptera: Aleyrodidae)

The two most damaging biotypes of Bemisia tabaci, B and Q, are sympatric in the Mediterranean basin and show high resistance to pyrethroids synergized by organophosphates. Previous work showed that in the B biotype, this resistance is associated with the L925I mutation in the para-type voltage gated sodium channel. Here we identified two mutations in the para-type voltage gated sodium channel associated with resistance to pyrethroids synergized by organophosphates in the Q biotype: the L925I mutation that occurs in the B biotype, and substitution of threonine to valine, at position 929 (T929V). To determine if the L925I and T929V mutations have single or multiple origins, we sequenced the DNA regions flanking the mutations from 13 B and Q strains collected worldwide. The survey identified five resistant alleles and five susceptible alleles. In the resistant alleles, the nucleotide diversity was low within biotypes (0.001), but high between biotypes (0.033). Nucleotide diversity in susceptible alleles was high between the two biotypes (0.028). These observations are consistent with multiple independent origins of resistance. Although the B and Q biotypes coexist in several regions of the Mediterranean basin, divergence in their DNA sequences at the para-type voltage gated sodium channel locus suggests gene flow between these biotypes is low or nil.     

Asymmetry in thermal tolerance trade-offs between the B and Q sibling species of Bemisia tabaci (Hemiptera: Aleyrodidae)

We investigated life history trade-offs related to thermal tolerance in two sibling species, commonly referred to as the B and Q biotypes, of Bemisia tabaci. We focused on basal resistance to short unpredicted heat stress, which reflects the organism investment, during both optimal and stressful conditions, in insuring its survival. At 27 °C, the relative reproductive performance of B was seven-fold higher than Q. After short stress of 42 °C, these differences increased to 23-fold. A turnover took place after short stress of 43 and 45 °C, in which Q adults performed better. As the expression of the analysed Hsp70 and other stress-related genes was found to be higher in the Q species, our data likely reflects two different strategies for optimal performance. B lowers soma protection for achieving maximum reproduction ('direct inhibitory' trade-off model), whereas Q invests significant resources in being always 'ready' for a challenge.     

Genetic diversity of Iberian populations of Bemisia tabaci (Hemiptera: Aleyrodidae) based on random amplified polymorphic DNA-polymerase chain reaction

The genetic structure of six Iberian populations of the whitefly Bemisia tabaci, two of them biotype Q, one biotype B, and the other three a mixture of both, has been studied using random amplified polymorphic DNA-polymerase chain reaction (RAPD-PCR). A total of 336 individuals was completely discriminated by means of 234 scored bands. Separate analyses of molecular variance of haploid males and diploid females using the pairwise number of differences between haplotypes showed that biotypes contribute significantly more to the observed variability than populations within biotypes. On average, gene flow between two biotypes of the same population is lower than between populations of identical biotypes. On the basis of these results and the nondetection under natural conditions of a single hybrid, we consider that both biotypes are genetically isolated under the ecological conditions prevailing in the south Iberian Peninsula. All populations of biotype Q presented similar values of intrapopulational diversity, which were higher than the values shown by populations of biotype B.