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.     

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.     

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.     

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.     

Performance of Myzus persicae (Hemiptera: Aphididae) clones on different host-plants and their host preference

The performance of eighteen clones of Myzus persicae (Sulzer) on pepper and tobacco plants at 20 degrees C and L16:D8 and the choice of young adult apterae between tobacco and pepper leaf-discs were examined. The clones were collected from weeds and peach in two tobacco-growing regions: Katerini, northern Greece and Karditsa, central Greece (only from weeds) and from Lehonia, central eastern Greece where tobacco is not cultivated. All clones did well on both hosts. However, the analysis of data revealed a significant effect of "region / host plant origin" on aphid performance. The mean values of adult weight, intrinsic rate of increase and fecundity of the clones collected in Lehonia and reared on tobacco were significantly lower than the observed values for clones from Katerini and Karditsa. Aphids from Lehonia had significantly higher mean values for developmental time on tobacco than clones from the other regions whereas the opposite was observed when aphids were reared on pepper. Aphids collected in Lehonia performed better on pepper than those originating from the tobacco-growing regions. A choice test revealed differences among the clones originating from different regions. Fifty three percent and 43% of aphids from weeds and peach from Lehonia, respectively, chose pepper. By comparison 41.5% and 40.0% of aphids from peach and weeds from Katerini, respectively and 49.5% of aphids from Karditsa preferred tobacco. The results are discussed in relation to host specialization in M. persicae.     

蚜虫对杨属植物不同寄生部位的形态适应(英文)

为了明确取食相同植物不同部位和营不同虫瘿的蚜虫是否存在形态适应,论文以取食杨属Populus植物的10种蚜虫为研究对象,基于蚜虫喙末端、各足跗节和爪的形态测量数据,对122个克隆的形态变异通过一般的统计描述、典型变量分析(canonical variates analysis,CVA)以及聚类分析等方法进行了研究。结果显示,不同的蚜虫克隆间形态特征存在分异,与不同的虫瘿类型和取食部位相关。3个典型变量分析明显地区分了形成虫瘿和不形成虫瘿的蚜虫克隆,在叶片上形成真虫瘿和伪虫瘿的克隆以及不产生虫瘿的克隆,并形成明显的克隆簇。而用于分析的形态学特征,如喙末端、跗节和爪的测量值,其一般统计描述的差异也支持这些区分。蚜虫自然种群不同克隆簇的区分很好地对应了不同的取食部位和不同的虫瘿类型,且各簇的形态特征体现了各自的特点,表明了蚜虫对杨属植物已经形成了良好的形态适应。同时,也初步讨论了不同蚜虫克隆簇形态适应产生的原因。并建议在基于形态特征探讨昆虫的系统发育关系和进行传统分类时,必须考虑昆虫形态特征的适应性;在深入研究昆虫与寄主植物相互之间关系时,需要将各种形态特征综合考虑并关注其它的影响因素。     

Separation of Myzus (Nectarosiphon) antirrhinii (Macchiati) from Myzus (N.) persicae (Sulzer) and related species in Europe (Homoptera: Aphididae)

Abstract. Multivariate morphometric analysis (canonical variates) was used to discriminate between closely related taxa within the Myzus persicae group. It is demonstrated that dark green, anholocyclic populations with a 2n=13 or 2n=14 karyotype in Europe, hitherto treated as a form or biotype of M.persicae, all conform to a discrete morphometric grouping and should therefore be treated as a separate taxonomic entity, permanently isolated from the M.persicae gene pool. It is suggested that this taxon was first described as Siphonophora antirrhinii by Macchiati in 1883. A discriminant function is provided to separate most individual apterae of antirrhinii from those of persicae. Myzus icelandicus Blackman sp.n., on Caryophyllaceae and other plants in Iceland, is distinguished from M.polaris H.R.L. and M.certus (Walker), and a key is given to apterous virginoparae of the species of the M.persicae group in Europe.     

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.     

Analysis of clonal diversity of the peach–potato aphid, Myzus persicae (Sulzer), in Scotland, UK and evidence for the existence of a predominant clone

Clones of the peach–potato aphid, Myzus persicae (Sulzer), mostly from Scotland, UK were examined using an rDNA fingerprinting technique. Eighty patterns (genotypes) were found amongst the 276 clones. A large number of clones (30%) from all sample areas in Scotland exhibited the same simple pattern, suggesting the presence of a single M. persicae clone. There was no difference in genotype distributions between M. persicae collected from brassica or potato crops, suggesting that host-adapted genotypes have no advantage in the field. Different fingerprints were randomly distributed in the environment, although clones taken from the same leaf were more often the same fingerprint. Highly distinctive fingerprints, which were more widely distributed, suggest that this technique could be used to follow individual clones. In addition to the common clonal type, multiple fingerprint bands were found over successive years, implying that, in Scotland, local overwintering asexual populations are the most common source of M. persicae in the following year.     

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.     

Morphometric correlates of karyotype and host plant in genus Euceraphis (Hemiptera: Aphididae)

Abstract Samples of birch- and alder-feeding aphids of genus Euceraphis were measured and analysed using multiple discriminant analysis (canonical variates) to find out if morphological variation correlated with previously reported differences of karyotype and host association. The dataset comprised groups of specimens defined solely by locality and collection date. Mean scores on the first two canonical variates clustered the samples fully in accordance with their karyotypes and host plants, confirming the existence of a number of morphologically similar but distinct host-specific taxa within the E. betulae group. Three new species are described: Euceraphis borealis Blackman, sp.n. on Betula glandulosa and B. nana , Euceraphis papyrifericola Blackman, sp.n. on B. papyrifera , and Euceraphis quednaui Blackman, sp.n. on B. occidentalis . A key is provided to alate viviparae of the genus.     

Morphometric analysis of Alaskan members of the genus Potentilla sect. Niveae (Rosaceae)

A morphometric study of Potentilla nivea, P unijlora, and P hookeriana, as well as the close relative of the latter, P furcata, has been carried out, and the quantitative data subjected to Canonical Discriminant Analysis. The four taxa belong to the arctic-alpine section Niveae of Potentilla, and material for the analysis was collected in Alaska, U. S. A. The a priori defined groups are based on petiole hair type, the qualitative, and only, character traditionally used to distinguish taxa within Potentilla sect. Niveae. The hair types recognized previously by taxonomists have been vaguely defined, and the intraspecific variation of other morphological characters has never been discussed. Ordination by canonical discriminant analysis was performed to characterize mean differences among species, to obtain insight into group differences, and to estimate character weights from correlations between canonical variates and original variables. The four taxa differ significantly in the canonical analysis of six quantitative characters. Leaflet length, incision depth (length of leaflet teeth), and ovule number are shown to be the most important discriminators. A key to the four taxa, taking into account the intra- vesus interspecific variation, as well as character weights, is provided.     

Microsatellite variation in cyclically parthenogenetic populations of Myzus persicae in south-eastern Australia

We examined the population structure of the introduced aphid, Myzus persicae collected mainly from its primary host, Prunus persica, in south-east Australia. Myzus persicae has been present in Australia since at least 1893. Samples were collected in the spring of 1998 from two mainland and three Tasmanian localities and isofemale lines were established in the laboratory. The reproductive mode (life cycle), karyotype and 17-locus microsatellite genotype of each clone were determined. All populations showed significant population differentiation (F(ST) 0.058-0.202) even over small geographic distances (<50 km). All clones were karyotypically normal except for a subset of clones from one site that was exposed to the carbamate insecticide, Pirimor, the week prior to sampling. Those clones were heterozygous for an autosomal 1,3 translocation frequently associated in M. persicae with insecticide resistance. In contrast to other loci and despite being on different chromosomes, loci myz2(A) and M55(A) showed general and significant linkage disequilibrium. These loci may be affected by epistatic selection. We discuss the observed high clonal diversity, moderate but significant population differentiation, general conformance to Hardy-Weinberg equilibria and low linkage disequilibria with particular focus on the global population biology of M. persicae.     

Size and shape in similarity coefficients based on metric characters

A variety of quantitative measures of distance and similarity have been used in physical anthropology, often without sufficient prior testing of their usefulness and understanding of their meaning. A variety of such coefficients were compared by applying them to an odontometric sample. It was found that Penrose's size-shape dichotomy extends not only to his coefficients but to all other similarity measures as well, such as the coefficient of racial likeness, generalized distance, canonical variates, and Q-mode correlation. Only size differences were detected by the C.R.L., Penrose's size distance, D², and canonical variates, and as a result these methods failed to produce an accurate classification. Penrose's shape distance and Q-mode correlation coefficients produced better results due to their determination of similarity on the basis of more important shape and morphological differences. The D² and canonical variates methods were converted to shape measures through Q-mode standardization of the raw data, whereupon they also produced more meaningful results.     

Climate in relation to deterioration of stored grain

Summary A hypothesis elucidating the basic relationship between abiotic climatic variates and the deterioration of stored grain by biotic agents was developed for the 41 crop districts in the Prairie Provinces of Canada. Principal component (7 variates measured) and canonical correlation (6 variates measured) analyses were applied to data from each of the 3 crop years (1967–1969), for each of the 3 principal component analyses and for each of the 3 canonical correlation analyses. The data consisted of crop district means of infestation reports from 2522 managers of grain elevators and also of representative meteorological observations from each crop district. The major variates measured were the number of reports of: hot spots, infestations of a) fungus beetles and mites, and b) Cryptolestes ferrugineus (Stephens), and grain fumigants. The climatic variates consisted of mean temperature, precipitation, and number of days above 20° C (68° F) for July, August and September. Three major components, which together explained 64–80% of the variation, indicated broadly similar trends in all analyses. The first component indicated that lower mean temperatures and larger numbers of cool days increased the incidence of fungus-induced hot spots in wet harvested grain. The second component indicated the extent of arthropod infestations: warmer weather increased C. ferrugineus infestation, whereas cooler weather increased infestation by mites and secondary insect species. The third component was a measure of precipitation. Canonical correlation analyses based on 1967 and 1969 data yielded canonical roots that were statistically insignificant at the 1% level. Analyses of 3 external abiotic and 3 biotic response variates based on 1968 data showed that only the first pair of canonical variates was significant, the degree of predictability (R _c ² ) being 56% (P<0.01). The analyses revealed that lower temperatures, a greater number of cool days, and higher precipitation increased the number of hot spots in 1968 confirming the findings of the principal component analyses. The pattern of hot spot development in relation to climatic characteristics was similar in all crop districts for all years examined. There was a linear relationship between climatic and biotic variates for all these crop districts but the degree of relationship varied geographically. The first and second principal component values for each crop district were ranked and plotted on a climatic map of the Prairie Provinces. The ranking showed that the incidence of hot spots and arthropod infestations was highest in the Sub-boreal and Humid southeastern regions, whereas the incidence was lowest in the Dry Belt region. C. ferrugineus was most common in the Humid southeastern region.Contribution No. 522 from Canada Department of Agriculture, Research Station, Winnipeg, Manitoba, R3T 2M9, Canada.     

Canonical correlation analysis of marine macrobenthos survey data

Canonical correlation analysis is applied to measurements of environmental variables and species distributions made during a survey of macrobenthos around a sewage-treatment farm drain. The implications of data reduction, necessary to enable the method to proceed, are discussed. The amount of data was reduced by discarding the rarest species, discarding species occurring at fewest stations, and including only those species and environmental variables which correlated highly with the greatest number of other variables. Only the third data-reduction scheme gave ecologically sensible results. Use of station scores on the first two canonical variates (CV1 and CV2) enabled the sampling grid to be divided into a group of nearshore stations, a group of intermediate depth, and a group of deep offshore stations. Loadings of environmental variables on the canonical variates were found to be unstable but correlations between these variables and canonical variates enabled the variates to be interpreted: CV1 as a gradient of depth and associated changes in sediment characteristics, CV2 with depth- and nutrient-related components, and CV3 as patchiness in sediment characteristics different from that normally expected with depth. Use of correlations between species and canonical variates enables definition of two major species groups, one confined to nearshore environments and a second offshore. These groups (and their sub-groups) related well to groups defined previously by hierarchical classification. It is concluded that, with careful attention to the method of data reduction, canonical correlation analysis can be an effective tool in the analysis of marine benthic survey data.     

The attack of the clones: tracking the movement of insecticide-resistant peach-potato aphids Myzus persicae (Hemiptera: Aphididae)

Myzus persicae (Sulzer) collected in Scotland were characterized for four microsatellite loci, intergenic spacer fingerprints and the resistance mechanisms modified acetylcholinesterase (MACE), overproduced carboxylesterase and knockdown resistance (kdr). Microsatellite polymorphisms were used to define a limited number of clones that were either fully susceptible to insecticides or possessed characteristic combinations of resistance mechanisms. Within these clones, intergenic spacer fingerprints could either be very consistent or variable, with the latter indicating ongoing evolution within lineages, most likely derived from the same zygote. Two clones (termed A and B) possessed all three resistance mechanisms and predominated at sites treated with insecticides. Their appearance on seed potatoes and oilseed rape in Scotland in 2001 coincided with extensive insecticide use and severe control failures. Clones C, I and J, with no or fewer resistance mechanisms, were found in samples from 1995 and were dominant at untreated sites in 2001. A comparison of Scottish collections with those from other UK and non-UK sites provides insight into the likely origins, distribution and dynamics of M. persicae clones in a region where asexual (anholocyclic) reproduction predominates, but is vulnerable to migration by novel genotypes from areas of Europe where sexual (holocyclic) reproduction occurs.     

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.