Co-existence of different host-adapted forms of the Myzus persicae group (Hemiptera: Aphididae) in southern Italy

Multivariate morphometric analysis (method of canonical variates) was used to compare 38 parthenogenetic lineages and three field collections of aphids of the Myzus persicae (Sulzer) group sampled on peach and tobacco in the Caserta region of southern Italy. Comparisons were also made with the morphology of lineages collected on peach in Lehonia, in central eastern Greece away from tobacco-growing regions, and on tobacco in Naphplion, in southern Greece. The lineages were measured after parthenogenetic rearing for one to three generations on the same host (potato) under constant conditions. As in previous work, the multivariate morphometric approach separated the aphids from peach from those feeding on tobacco. The life cycle category of the lineages was also examined. Almost all the lineages from peach in Caserta were holocyclic, yet the lineages from tobacco in the same region were unable to produce sexual morphs. The results suggest that tobacco-adapted and non-tobacco-adapted forms co-exist in the same region in southern Italy, not only because they colonize different host plants, but also because they have different life cycles. This is in complete contrast to the situation in peach-growing areas of northern Greece, and shows that the ecology and population structure of M. persicae is different in neighbouring counties of the Mediterranean area, even where climatic conditions and cultivated crops are similar.     

Host-correlated morphological variation of Myzus persicae (Hemiptera: Aphididae) populations in Greece

Morphological variation in nine characters of 157 clones of Myzus persicae (Sulzer) was examined by multivariate analysis. The clones were collected from peach, Prunus persica, the primary host and the secondary hosts tobacco, Nicotiana tabacum, cabbage, Brassica oleracea, sugarbeet, Beta vulgaris and pepper Capsicum annuum. The 156 clones originated from various regions of Greece, both in the north, where a large part of the population has an annual bisexual generation on peach, and in more southerly regions, where populations are predominantly unisexual. One clone was collected from tobacco in Caserta, Italy. All clones were laboratory-reared on potato. Canonical variate analysis, hierarchical cluster analysis and a non-parametric classification tree method both revealed morphological differences associated with the host-plant on which they were collected. The scores of the first two canonical variates separated the tobacco-feeding clones from those originating from other secondary host-plants. However, in tobacco-growing areas the tobacco-feeding form predominated in spring populations on peach, and was sometimes found on other secondary hosts. In addition, using cluster analysis, the clones from tobacco which were sampled in the most southeasterly region showed a relatively large phenotypic distance from those collected further north and west. Moreover, clonal phenotypes were affected both by host plant and by long-term parthenogenetic rearing. However, in spite of these effects, the tobacco form was generally distinguishable from aphids originating from other hosts, indicating that the difference must have a genetic basis. In separate analyses of the clones originating from secondary hosts no association was found between morphology and either life cycle category or colour. Discriminant analysis showed that 89% of 1723 specimens could be correctly classified into the two groups.     

Genetic variation in Myzus persicae populations associated with host-plant and life cycle category

Random amplified polymorphic DNA (RAPD) analysis was applied on 96 clones of Myzus persicae (Sulzer) (Homoptera: Aphididae) representing seven populations collected from different host-plants and regions of Greece. Ten decamer random primers were used to evaluate genetic variation among the examined samples. Despite the variability found between clones, no specific RAPD marker was detected to discriminate the different populations. A significant finding was that aphids from peach and pepper, which were collected far away from tobacco-growing regions, especially those from peach, showed genetic divergence from the tobacco-feeding clones. Moreover, data analysis revealed a significant genetic divergence between holocyclic and anholocyclic populations from tobacco. Lastly, holocyclic clones showed higher level of estimated heterozygosity than the nonholocyclic (anholocyclic, androcyclic and intermediate) ones.     

Variation in the life cycle and morphology of the tobacco host-race of Myzus persicae (Hemiptera: Aphididae) in relation to its geographical distribution

Morphological variation and life cycle category were examined in 121 clones of Myzus persicae (Sulzer). The clones were collected from tobacco from three localities in Greece (Xanthi, Nea Efessos and Naphplion), one in Germany (Rheinstetten), one in France (Bergerac) and one in Spain (Madrid). Before morphometrics, all aphids were laboratory-reared on potato. The morphological variation was investigated using both canonical variates analysis and a novel non-parametric classification tree method. The life cycle category was examined by rearing the clones for three generations under short day conditions. In Nea Efessos a relative high proportion of clones was found to overwinter as eggs on the primary host. In the other regions all collected clones were non-holocyclic. Intermediate genotypes were found in all regions at percentages ranging from 4.0 to 24.0%. Androcyclic clones were found only in Xanthi, Greece (4.0%) and Rheinstetten, Germany (16.7%). The canonical variates analysis and the tree classification method revealed important intrapopulation polymorphisms in clones from Bergerac, Nea Efessos and Madrid. Both methods separated the populations originating from Greece from those collected elsewhere in western Europe. The observed morphological variation was probably due to genetic differences, since all clones were reared in a common environment. The results are discussed in relation to factors responsible for genetic divergence in M. persicae populations.     

Life cycle variation of Myzus persicae (Hemiptera: Aphididae) in Greece

During the years 1995-1999 the life cycle category of 2797 clones of Myzus persicae (Sulzer) was examined. The clones originated from primary and secondary hosts from different localities of North and Central Greece and the island of Crete in the south. Four different overwintering life cycle strategies were found that have also been described for M. persicae and other heteroecious species previously. A geographical variation was found in the proportion of holocyclic clones from tobacco and other secondary hosts associated with the abundance of the primary host in the sampling regions. In Central Macedonia, around the main peach-growing regions, the proportion of holocyclic clones was mostly above 50% and in some cases reached 100%. In localities of East Macedonia, holocyclic clones were also frequent. On the other hand, further south or in north-eastern Greece, where peach is not common, the proportion of holocyclic clones varied between 0 and 33%. Fifty seven percent of examined anholocyclic clones produced males under short day conditions, suggesting that androcyclic clones in Greece represent an important factor of genetic variability. Intermediate clones were sampled from all host-plants but at low frequencies (3.6% of total examined clones and 6.9% of non-holocyclic ones). Moreover, a regional variation was found in different colour forms feeding on tobacco plants. Red clones were predominant in regions where aphids overwinter parthenogenetically on weeds or winter crops. However, almost all clones from the primary host were green. The ecological aspects of life cycle variation are discussed.     

An interval timer controls the production of sexual morphs in Myzus persicae (Homoptera: Aphididae)

Abstract The interval timer mechanism was examined in two clones of Myzus persicae (Sulzer) originating from peach in Naousa, North Greece. Overwintering eggs were collected in April 1997 and after egg hatch the morphs of progeny were recorded at LD 10 : 14 h and 17 °C until the appearance of sexuals. In one clone, observations were continued until the 21st generation after egg hatch. In addition, the production of sexual morphs was examined in two other clones originating from Lehonia, Central Greece, when apterae (wingless females) were transferred from long to short days and 17 °C in the second and eighth generation of parthenogenetic rearing after their collection from peach trees. In clones from Naousa, the first gynoparae appear and comprise 10% of alate (winged) females in the second half of the third generation after egg hatch. The percentage of gynoparae increased in each consecutive generation, reaching 100% in fifth and seventh generations in clones Nb and Na, respectively. First males appeared later than gynoparae, in fifth and sixth generation in clones Nb and Na, respectively. In clone Nb, the number of males, after their appearance in fifth generation, was relatively constant from the sixth to the ninth generation (10.8–12.3). Then a significant increase was observed in the 10th generation, thereafter remaining relatively constant until the 21st generation (18.1–19.8). A similar trend in the number of males produced was observed in clones deriving from Lehonia.     

Distribution of common genotypes of Myzus persicae (Hemiptera: Aphididae) in Greece, in relation to life cycle and host plant

Microsatellite genotyping was used to identify common clones in populations of the Myzus persicae group from various hosts and regions in mainland Greece and southern Italy and to compare their distribution and occurrence on tobacco and other crops. Common clones were defined as genotypes collected at more than one time or in more than one population; and, therefore, unlikely to be participating in the annual sexual phase on peach. Sixteen common genotypes were found, accounting for 49.0% of the 482 clonal lineages examined. Eight of these genotypes were subjected, in the laboratory, to short days and found to continue parthenogenetic reproduction, i.e. they were anholocyclic. Four of the six commonest genotypes were red, and one of these accounted for 29.6% of the samples from tobacco and 29.4% of those from overwintering populations on weeds. All six commonest genotypes were found on weeds and five of them both on tobacco and on other field crops. In mainland Greece, the distribution of common clones corresponded closely with that of anholocyclic lineages reported in a previous study of life cycle variation. Common genotypes were in the minority in the commercial peach-growing areas in the north, except on weeds in winter and in tobacco seedbeds in early spring, but predominated further south, away from peach trees. This contrasts with the situation in southern Italy, reported in a previous paper, where peaches were available for the sexual phase, yet all samples from tobacco were of common genotypes.     

Intraspecific variation in zucchini yellow mosaic virus transmission by Myzus persicae and the impact of aphid host plant

Three isofemale lines of Myzus persicae (Sulzer), two lines collected from and reared on a brassicaceous host, and one line collected from and reared on a malvaceous host, were evaluated for their efficiency of transmitting Zucchini yellow mosaic virus (family Potyviridae, genus Potyvirus, ZYMV). In the first experiment, the transmission efficiencies of two clones from Brassicaceae (B1 and B2) were 52.0 and 60.8%, respectively, and these transmissions were not significantly different. In a second experiment, the transmission efficiencies of the clone on Malvaceae (M1) and clone B2 were significantly different at 35.6 and 55.7%, respectively. Further experiments evaluated host-related mechanisms that may have contributed to the differential transmissions observed between clones M1 and B2. Studies on short-term feeding showed that aphids continuously reared on okra, Abelmoschus esculentus (L.) Moench (malvaceous host), and those that were reared on okra and allowed a 24-h preacquisition feeding period on mustard, Brassica juncea (L.) Czern (brassicaceous host), had significantly lower transmission than aphids continuously maintained on mustard. Aphids reared on mustard and allowed a 24-h preacquisition feeding period on okra had intermediate transmission efficiency. In long-term host association studies, we found that aphids reared on mustard had significantly higher transmission efficiency than those reared on okra, and aphids reared first on okra and then switched to mustard had a transmission efficiency that was intermediate and not significantly different from the other two treatments. Our study reveals the existence of intraspecific variation in the transmission of ZYMV by M. persicae, and it suggests that to accurately assess the transmission capability of ZYMV by this species, multiple clones should be examined. Furthermore, the host plant on which the aphid is reared as well as the host plant on which it feeds just before virus acquisition contribute to ZYMV transmission efficiency of M. persicae.     

3种寄主上桃蚜的选择性及形态分化

桃蚜是一种重要的农业害虫,寄主广泛,种下分化复杂。以采自黄土高原旱作区桃树、烟草、甘蓝上的桃蚜为研究对象,通过叶片选择法、传统比较形态测定法研究了3种寄主上桃蚜的选择性及形态分化。结果表明:在3种寄主同时存在的情况下,烟草上的桃蚜嗜食烟草,表现为63.5%的桃蚜选择烟草叶,13.8%选择甘蓝叶,8.2%选择桃叶,而甘蓝和桃树上的桃蚜对原寄主没有表现出明显的嗜好性;从形态指标来看,3种寄主上的桃蚜在体长、触角末节长度、后足腿节长度、触角与体长的比例方面存在显著差异(P0.05),说明这几个特征可以作为区分这3种寄主上桃蚜的依据。综合分析可以初步认为黄土高原旱作区烟草上的桃蚜可能形成了寄主专化型-烟草型。     

Production of sexual morphs by apterous virginoparae of Myzus persicae (Hemiptera: Aphididae) in relation to pre- and postnatal exposure to short day conditions

The production of males and females by apterae of a holocyclic clone of Myzus persicae (Sulzer) was studied in the laboratory by transferring apterous virginoparae from long day to short day conditions at different stages of their pre- and post-natal development. Prenatal exposure was also examined in nine additional clones coming from different regions of Greece. By increasing the number of short day cycles at 17 degrees C, the aphids switched to male production earlier and produced more males. The highest mean number of male progeny was observed in apterae that were transferred to short day conditions 13-14 days prenatally. Apterae that were transferred to short day conditions after the fourth nymphal instar produced only females. Apterous virginoparae were produced by aphids that were transferred to short day conditions 0-1 days prenatally or postnatally. Aphids, given short days prenatally or in the first instar, produced alate female progeny that were all gynoparae. The later the instar aphids were transferred to short day conditions the fewer the gynoparae produced among their female progeny. A reduction in the total number of progeny and embryos per ovariole was observed as aphids were transferred to short day conditions in earlier stages of their development. The increase in male production, which follows the increase in the duration of long night exposure, was also observed in the additional eight out of nine examined clones.     

Evaluation of artificial diets for rearing Aphis glycines (Hemiptera: Aphididae)

Artificial aphid diets have been previously developed for the pea aphid, Acyrthosiphon pisum (Harris), and the green peach aphid, Myzus persicae (Sulzer). The ability to rear aphids on an artificial diet allows for selectively adding or subtracting compounds from an aphid's food source to determine the effect on fecundity and longevity. Five diets previously developed for the green peach aphid and the pea aphid were tested for their suitability for rearing soybean aphid, Aphis glycines Matsumura. The best diet, originally developed for the green peach aphid and based on the amino acid profile of young potato plants, allowed 12 generations of soybean aphids to develop. For all diets tested, aphid fecundity, and longevity were greatly reduced in comparison with aphids reared on soybean, Glycine max (L.) Merr., plants or on detached soybean leaves. In addition, mean developmental time was significantly longer for aphids reared on artificial diets.     

Recent changes in the distribution of carboxylesterase genes and associated chromosomal rearrangements in Greek populations of the tobacco aphid Myzus persicae nicotianae

We present data on the frequency of amplified E4 and FE4 carboxylesterase genes in Myzus persicae s.l. clones collected during the years 2002–2007 and 2012 in Greece. Most clones were of the tobacco aphid, Myzus persicae nicotianae. Samples from 2012 were genotyped with microsatellite DNA markers and a number of them were karyotyped. Aphid clones with amplified FE4 genes predominated in all years, whereas E4 was present in only 3.5% of all samples and always occurred in clones with FE4. Most of the clones examined showed high carboxylesterase activity levels (R2 resistant category). The results showed marked changes in the frequencies of the two carboxylesterase genes in the tobacco aphid populations compared to published data that were collected in Greece in the mid 1990s, when E4 was recorded on its own in 20% of all samples and in 32% of samples from tobacco. A parallel change in karyotype was also observed because the A1,3 translocation, which had a worldwide association with amplified E4 genes in the 1990s, was not detected in the clones analyzed in 2012. Possible causes for these changes are discussed, although selection as a result of pest management practices appears to be the major one. Novel chromosomal rearrangements were also found in M. persicae nicotianae clones. These rearrangements could be a result of clastogenic effects of nicotine, which could persist because of the holocentric nature of aphid chromosomes. The results are discussed in relation to rapid evolution events that have taken place in the tobacco aphid in Greece during the last two decades. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 455–470.     

Host choice by Aphidius colemani: effects of plants, plant–aphid combinations and the presence of intra-guild predators

Abstract.  1. This study first measured the effect of plant (radish, pepper, and wheat), aphid–plant combination (the green peach aphid Myzus persicae on pepper and radish, and the bird cherry-oat aphid Rhopalosiphum padi on wheat) and the host on which Aphidius colemani was reared (the cotton aphid Aphis gossypii on cucumber, M. persicae on radish and pepper, and R. padi on wheat) on host choice behaviour of the parasitoid and the performance of its offspring. Then, the effect of predator presence ( Coccinella undecimpunctata larvae) on host preference of the wasps was tested.
2. When reared on M. persicae on either radish or pepper , wasps preferred the aphid–plant combination from which they had emerged. Wasps reared on A. gossypii (naïve to all hosts offered) and R. padi preferred to parasitise M. persicae on radish and M. persicae on either radish or pepper, respectively. Rhopalosiphum padi on wheat was the least preferred and also the least suitable host, as determined by offspring body size.
3. Contrary to expectations, the presence of predators did not influence the host choice of A. colemani , even when predator-free plants were offered nearby. Results indicate that wasps are more likely to remain in some host systems but not in others, even when facing risk of predation.
4. Results are discussed in respect to field data and the relative importance of risk of predation and host preference to wasp fitness.     

Survival, growth, and reproduction of two aphid species on sucrose solutions containing host or non-host honeydews

Aphids, like most phloem-feeding insects, commonly exhibit a high degree of host specificity. Plant-specific chemical compounds are likely to serve as important host selection cues for monophagous aphids and such substances could be present in aphid honeydew. Apterous virginoparae ofMyzus persicae (Sulzer) andPhorodon humuli (Schrank) were reared on a buffered sucrose solution containing various aphid honeydews or a mixture of amino acids. In two separate experiments, the host-specificP. humuli (hop aphid) could grow and reproduce only on diets containing honeydew collected from hop (Humulus lupulus L.).M. persicae (the green peach aphid, GPA) did not perform well on diets containing hop honeydew, perhaps because hop is a poor GPA host. Honeydew collected from preferred GPA host plants rape,Brassica napus L., and jimsonweed,Datura stramonium L., allowed GPA growth and reproduction. Hop aphids, however, performed poorly on rape and jimsonweed honeydew diets. Bell pepper,Capsicum annuum L., honeydew supported neither the hop aphid nor GPA. The study of aphid honeydew components may contribute towards a more complete understanding of host preference and selection phenomena in aphids.     

Managing Brachycaudis persicae (Homoptera: Aphididae) during peach orchard establishment

We investigated whether preplant root dips or postplant sprays or drenches could be used to control the black peach aphid, Brachycaudus persicae (Passerini), on young peach trees. In greenhouse trials, B. persicae died after being placed on peach trees that had their roots dipped in imidacloprid before planting. All rates (0.8, 1.6, and 3.2 ml/liter of water) tested controlled this aphid. B. persicae died faster on trees dipped in imidacloprid (1.6 ml/liter of water) for 5 min before planting than on trees dipped for 2 s. Aphids eventually died on all treated trees. Approximately 50% of untreated trees died from B. persicae infestations in greenhouse studies. Results from afield experiment show that peach trees root-dipped in imidacloprid before planting or drenched with imidacloprid after planting eliminate B. persicae infestations and prevent root colonization for at least 1 yr.     

Movement, toxicity, and persistence of imidacloprid in seedling Tabasco pepper infested with Myzus persicae (Hemiptera: Aphididae)

Application of imidacloprid to the soil in which Tabasco pepper, Capsicum frutescens L., seedlings were growing was highly effective against the green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae). In just 48 h after the soil drench, aphid numbers on treated plants declined from 292.1 to 33.0 per plant, a reduction of 89%. By 72 and 96 h after the application, the reductions were 97 and 100%, respectively. Reductions in green peach aphid numbers also indicated that imidacloprid readily moved throughout the Tabasco pepper plant. Although, initial green peach aphid reductions at 24 and 48 h after imidacloprid application to soil, were greater on the lower leaves than on the upper leaves, by 72 h toxicity was high throughout the plant. At 48 h, overall green peach aphid reduction on seedlings grown in wet soil was significantly higher than that on plants growing in the drier soil. Regardless of soil moisture or leaf location, no live green peach aphids were detected on treated seedlings after 96 h. After the initial uptake period, toxicity to green peach aphid remained high for 5 wk. Under Tabasco pepper production conditions in Central America, the greatest need for aphid management is just after transplanting. Imidacloprid soil drenches before transplanting should offer the Tabasco pepper producer an extended period of aphid-free production.     

桃蚜自然种群初级和次级共生菌的分子鉴定

蚜虫不仅带有专性初级内共生菌,还常常带有不同类群的兼性次级共生菌。本研究采用真细菌16S rDNA的通用引物,从桃蚜Myzus persicae (Sulzer)烟草种群体内扩增出1.5 kb的共迁移目的条带,然后将其克隆,并在阳性克隆筛选时进行了RFLP分析。当采用EcoRⅠ进行单酶切分析时,目的片段被切割成～650 bp和～850 bp两个小片段;当采用EcoRⅠ和HindⅢ进行双酶切分析时,该蚜群共生菌酶切谱带被分成2组,其中1组(GroupⅠ)只具有一个EcoRⅠ酶切位点,而另1组(Group Ⅱ)不仅具有一个EcoRⅠ酶切位点,还具有一个HindⅢ酶切位点,而且GroupⅠ为优势共生菌群。在此基础上分别对2组共生菌的16S rDNA全序列（～1.5 kb）进行了测定,结果表明：GroupⅠ属于泛菌属Pantoea,与成团泛菌Pantoea agglomerans亲缘关系最近（同源性达99.70%）,而Group Ⅱ属于蚜虫的专性内共生菌,即Buchnera aphidicola（同源性达99.50%）。这是在蚜虫体内存在泛菌次级共生菌的首次报道。     

MORPHOLOGICAL STUDIES ON MYZUS PERSICAE IN CHINA

Abstract  Experiments were carried out to study the morphological characters of Myzus persicae (Sulzer) on tobacco, rape and peach from different localities of China. The results showed that the number of accessory hairs on ultimate rostral segment had no difference among all samples in 14 morphological characters used by Blackman in this study. Cluster analysis indicated that the morphological differences between samples from peach and samples from other host plants was the biggest, the next was the samples from Changchun on tobacco. Compared with peach, the aphid from rape was more similar to that from tobacco in morphology. For the samples on tobacco from the central and southern China, they can be clustered into yellowish-green group and reddish-brown group. It shows unlikely to separate successfully the aphids on tobacco from those on non-tobacco host plants only by morphological characters.     

中国桃蚜形态学研究(英文)

对来自中国不同地区的烟草、油菜和桃树的桃蚜形态特征研究结果表明：在所研究的14个形态特征中,喙末节端部的次生毛的数目这一形态特征在所有的样本中均无显著差异。聚类分析表明采自桃树上的桃蚜的形态特征与采自其它寄生植物上的桃蚜的形态特征差异最大;其次是采自长春烟草上的桃蚜。油菜上的桃蚜在形态上比桃树上的桃蚜更接近烟草上的桃蚜。我国中南部烟草上的桃蚜又可区分为黄绿和红褐两类。在本研究中所用的14个形态特征不能明确地将我国烟草上的桃蚜与其它寄生植物上的桃蚜区分开。     

Antibiosis and antixenosis of six commonly produced potato cultivars to the green peach aphid, Myzus persicae Sulzer (Hemiptera: Aphididae)

The antibiotic and antixenotic resistance of six commonly produced potato cultivars in Iran including Aozonia, Agria, Cosima, Cosmos, Kondor and Savalan to the green peach aphid, Myzus persicae Sulzer, were investigated under laboratory conditions at 20 ± 2oC, 65 ± 5% RH and 16:8h (L:D) in 2009. Antibiosis experiments showed significant differences in the developmental time, nymphal survivorship, fecundity, adult longevity of the green peach aphid among the potato cultivars. Intrinsic rate of natural increase (r m) for apterous aphids varied significantly with the potato cultivars on which the aphids were reared. This value ranged from 0.225 to 0.293 females/female/day, which was lowest on Cosmos and highest on Aozonia. Additionally, the estimated net reproductive rate (R0) and finite rate of increase (λ) for apterous aphids were the lowest on Cosmos. For the antixenosis experiment, no significant difference was found in aphid's preference to the potato cultivars. However, Aozonia was preferred more than the other five cultivars by the apterous aphids. Therefore, our results demonstrated that among the investigated cultivars the Cosmos cultivar is moderately resistant to the green peach aphid.