Aphis （Hemiptera： Aphididae） species groups found in the Midwestern United States and their contribution to the phylogenetic knowledge of the genus

A phylogeny of the genus Aphis Linnaeus, 1 758 was built primarily from specimens collected in the Midwest of the United States. A data matrix was constructedwith 68 species and 41 morphological characters with respective character states of alate and apterous viviparous females. Dendrogram topologies of analyses performed usingUPGMA （Unweighted Pair Group Method with Arithmetic Mean）, Maximum Parsimony and Bayesian analysis of Cytochrome Oxidase I, Elongation Factor 1-α and primary endosymbiont Buchnera aphidicola 16S sequences were not congruent. Bayesian analysis strongly supported most terminal nodes of the phylogenetic trees. The phylogeny wasstrongly supported by EFI-α, and analysis of COl and EFI-α molecular data combined with morphological characters. It was not supported by single analysis of COI or Buch-hera aphidicola 16S. Results from the Bayesian phylogeny show 4 main species groups： asclepiadis,fabae, gossypii, and middletonii. Results place Aphis and species of the generaProtaphis Bōrner, 1952, Toxoptera Koch, 1856 and Xerobion Nevsky, 1928 in a monophyletic clade. Morphological characters support this monophyly as well. The phylogenyshows that the monophyletic clade of the North American middletonii species group belong to the genus Protaphis： P. debilicornis （Gillette ＆ Palmer, 1929）, comb. nov., P. echinaceae（Lagos and Voegtlin, 2009）, comb. nov., and P. middletonii （Thomas, 1879）. The genus Toxoptera should be considered a subgenus of Aphis （stat. nov.）. The analysis also indicatesthat the current genus Iowana Frison, 1954 should be considered a subgenus of Aphis （stat. nov.）.     

Presence of two host races of Aphis gossypii Glover (Hemiptera: Aphididae) collected in Turkey

The aphid, Aphis gossypii, is a primary pest of citrus, cotton, cucurbits and greenhouse‐grown vegetables in Turkey and throughout Europe. There is some previous empirical data suggesting that host‐adapted genotypes of this aphid exist which may in fact be host‐races. To determine if host races of A. gossypii are indeed present in the eastern Mediterranean region of Turkey, reciprocal host transfer experiments and life table analyses were performed with multiple asexual lineages (= clones) of the aphid collected from different hosts. The collection hosts included citrus, cucumber, eggplant, okra, sweet pepper and cotton. Aphid developmental times on the host from which the aphid was originally collected (= collection or natal host) were shorter (5.2–6.0 days) and had a higher intrinsic rate of population growth (r_m = 0.25–0.44) than the 6.6–7.3 days required when the aphid was reared on a non‐original collection host (= non‐collection host or non‐natal host) and had r_m = 0.03–0.30. Total immature mortality of the cotton clone, especially in the first nymphal stage, was high (51–100%) with low r_m (0–0.03) on cucumber, citrus and sweet pepper. Aphid populations transferred from citrus, eggplant and okra to cotton (r_m = 0.29–0.30) did not differ significantly in their performance from that of the cotton population on cotton (r_m = 0.34), whereas that from sweet pepper and cucumber populations (r_m = 0.22–0.24) were significantly lower. These data have allowed us to separate A. gossypii into two distinct biological groups: (a) a ‘generalist’ population obtained from cucumber, sweet pepper, citrus, eggplant and okra which exhibited statistically better development on cotton; versus (b) a population from cotton which, by comparison on reciprocal hosts, developed poorly on non‐natal hosts except on eggplant. Development of the cotton clone on cucumber and okra was not improved after four successive generations on the non‐natal host. The good development of A. gossypii from eggplant and cotton on these reciprocal hosts suggests that these particular clones were similar, if not identical, host races.     

Molecular phylogeny and divergence times of Hormaphidinae (Hemiptera: Aphididae) indicate Late Cretaceous tribal diversification

The aphid subfamily Hormaphidinae is a good candidate for the study of the evolution of insect – plant relationships. Most hormaphidine species depend on woody primary host plants and woody or herbaceous secondary host plants, and represent high host specificity, especially to their primary hosts. No detailed molecular phylogeny of Hormaphidinae has been reported, and the taxonomic positions of some taxa in this group remain unclear. To reconstruct major phylogenetic relationships and to understand the evolution of host association patterns for major lineages, we present the first detailed molecular phylogeny of Hormaphidinae, as inferred from nuclear and mitochondrial DNA sequences, using maximum parsimony, maximum likelihood, and Bayesian methods. The monophyly of Hormaphidinae and its three traditional tribes was supported, and a sister relationship between Hormaphidini and Nipponaphidini was suggested. Most inner relationships within tribes were also supported, and some novel relationships were revealed. Two subtribes of Cerataphidini are proposed. Divergence times estimated using a Bayesian approach indicate that tribal diversifications occurred during the Late Cretaceous and were coincident with the appearance of their primary host plants. The current pattern of secondary host association for the three tribes may have evolved in different time ranges. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 73–87.     

Molecular and morphological differentiation between Aphis gossypii Glover (Hemiptera,Aphididae) and related species,with particular reference to the North American Midwest

The cotton aphid, Aphis gossypii, is one of the most biologically diverse species of aphids; a polyphagous species in a family where most are host specialists. It is economically important and belongs to a group of closely related species that has challenged aphid taxonomy. The research presented here seeks to clarify the taxonomic relationships and status of species within the Aphid gossypii group in the North American Midwest. Sequences of the mitochondrial cytochrome oxidase 1 (COI), nuclear elongation factor 1-α (EF1-α), and nuclear sodium channel para-type (SCP) genes were used to differentiate between Aphid gossypii and related species. Aphis monardae, previously synonymised with Aphid gossypii, is re-established as a valid species. Phylogenetic analyses support the close relationship of members of the Aphid gossypii group native to North America (Aphid forbesi, Aphid monardae, Aphid oestlundi, Aphid rubifolii, and Aphid rubicola), Europe (Aphid nasturtii, Aphid urticata and Aphid sedi), and Asia (Aphid agrimoniae, Aphid clerodendri, Aphid glycines, Aphid gossypii, Aphid hypericiphaga, Aphid ichigicola, Aphid ichigo, Aphid sanguisorbicola, Aphid sumire and Aphid taraxicicola). The North American species most closely related to Aphid gossypii are Aphid monardae and Aphid oestlundi. The cosmopolitan Aphid gossypii and Aphid sedi identified in the USA are genetically very similar using COI and EF1-α sequences, but the SCP gene shows greater genetic distance between them. We present a discussion of the biological and morphological differentiation of these species.     

两种毛蚜线粒体基因组比较分析

【目的】线粒体基因组分析已被应用于昆虫系统发育研究。本研究以蚜科Aphididae重要类群毛蚜亚科物种为代表,测定并比较分析了该类蚜虫的线粒体基因组特征,探讨了基于线粒体基因组信息的蚜虫系统发育关系重建。【方法】以毛蚜亚科三角枫多态毛蚜Periphyllus acerihabitans Zhang和针茅小毛蚜Chaetosiphella stipae Hille Ris Lambers,1947为研究对象,利用长短PCR相结合的方法测定线粒体基因组的序列,分析了基因组的基本特征;基于在线t RNAscan-SE Search Server搜索方法预测了t RNA的二级结构;基于12个物种(本研究获得的2个物种和10个Gen Bank上下载的物种数据)的蛋白编码基因(PCGs)序列,利用最大似然法和贝叶斯法重建了蚜科的系统发育关系。【结果】两种毛蚜均获得了约94%的线粒体基因组数据,P.acerihabitans获得了14 908 bp,控制区为1 205 bp;C.stipae获得了13 893 bp,控制区为609 bp。两种毛蚜同时获得33个基因,包含接近完整的13个蛋白编码基因(PCGs)(nad5不完整),18个tRNA,2个rRNA基因;ka/ks值表明,C.stipae的进化速率更快。从基因组组成、基因排列顺序、核苷酸组成分析、密码子使用情况、t RNA二级结构等特征来分析,两种蚜虫线粒体基因组基本特征相似。系统发育重建结果表明毛蚜亚科、蚜亚科的单系性得到了支持,毛蚜亚科位于蚜科的基部位置。【结论】两种毛蚜线粒体基因组的基本特征相似,符合蚜虫线粒体基因组的一般特征,两种线粒体基因组的长度差异主要来自控制区长度的不同;系统发育重建支持毛蚜亚科与蚜亚科的单系性,毛蚜亚科位于蚜科较为基部的位置。研究结果为蚜虫类系统发育重建提供了参考。     

Aspects of insecticide resistance in the melon and cotton aphid, Aphis gossypii (Hemiptera: Aphididae)

Strains of the aphid Aphis gossypii, taken from cucumber in UK glasshouses were tested for resistance to pirimicarb, diazinon and heptenophos. Compared to a susceptible strain, resistance factors to diazinon of up to 659 were found in populations collected between 1987 and 1989. Compared to the same susceptible strain, diazinon-resistant populations also showed a reduced response to heptenophos. All field populations tested against pirimicarb were resistant. Host plant preference tests showed that all strains (including stock cultures) tested could readily breed on cotton. However strains originally collected from chrysanthemum did not transfer to cucumber and likewise strains from cucumber would not colonise chrysanthemum. Polyacrylamide gel electrophoresis of four of the aphid strains showed differences in migration rates and intensity of staining of esterase bands. Intensity of staining was correlated with the degree of diazinon resistance.     

多异瓢虫食物消耗与利用与其猎物棉蚜的寄主植物种类相关

【目的】猎物质量和类型在捕食者食物选择、消耗及营养转化过程中起着重要作用,植物的营养质量影响植食者的营养质量,进而植食者作为捕食者猎物的营养质量。为进一步了解植物-猎物-捕食者三营养层的相互作用,研究了寄主植物对捕食者食物消耗与利用的影响。【方法】在室内评价了多异瓢虫Hippodamia variegata (Goeze) 4龄幼虫和雌成虫取食和利用 5种不同葫芦科植物上的棉蚜 Aphis gossypii Glover的效率。【结果】多异瓢虫4龄幼虫和成虫均对南瓜Cucurbita moschata var. melonaeformis上的蚜虫取食量最高,而对瓢葫芦Lagenaria siceraria var. gourda上的蚜虫取食量最低。4龄幼虫取食哈密瓜 Cucumis melo var. cantalupensis上的蚜虫时,其相对生长率、食物利用率和食物转化率均达到最大,而雌成虫在取食黄瓜Cucumis sativus和搅瓜Cucurbita pepo var. medullosa上的蚜虫时,其相对生长率不存在显著差异,在取食黄瓜、瓢葫芦和搅瓜上的蚜虫时,其食物利用率和食物转化率也不存在显著差异。因此,哈密瓜上的瓜蚜更适宜作为多异瓢虫4龄幼虫的猎物,而黄瓜、瓢葫芦和搅瓜上的瓜蚜更适宜作为成虫的猎物。【结论】寄主植物种类与多异瓢虫对棉蚜的捕食效率相关,该结论为进一步利用昆虫的食性对害虫进行控制奠定理论基础。     

棉花型和黄瓜型棉蚜(Aphis gossypii Glover)的寄主适应性及转移通道

采用寄主转接建立生命表的方法研究了棉花型和黄瓜型棉蚜对不同寄主植物的适应性,以及两寄主型棉蚜是否可通过中间桥梁寄主实现寄主互换的问题。结果表明,两寄主型棉蚜直接互换寄主后,其存活和繁殖力显著下降,表现为棉花型和黄瓜型棉蚜的净增殖率比在原寄主上分别下降980倍和12倍,平均世代寿命缩短5～12d。两寄主型棉蚜均能利用木槿植物,并且适应性没有显著差异。但是两寄主型棉蚜均不能在车前草和大叶黄杨上存活和繁殖后代。西葫芦作物对棉蚜在木槿、棉花和黄瓜寄主上的相互转移起到了重要的桥梁寄主作用。冬寄主木槿上棉蚜可通过甜瓜或西葫芦转移到黄瓜寄主上,棉花和黄瓜上棉蚜也可通西葫芦作物分别转移到黄瓜和棉花作物上,从而形成棉蚜在不同寄主植物间的相互转移通道,造成为害和病毒病的扩张。     

Different exposure methods to neonicotinoids influenced biochemical characteristics in cotton aphid,Aphis gossypii Glover (Hemiptera:Aphididae)

We measured lipid, carbohydrate and protein content in three strains of cotton aphid, Aphis gossypii (Hemiptera: Aphididae) from very resistant to neonicotinoids up to semi sensitive and sensitive strains in different exposure method of spraying to neonicotinoids. We observed changes in energy source rates at which each substrate was metabolised under starvation, selection and residue stress method spraying that assist in metabolisation of their biochemical parameters. These particular exposure methods influenced some of biochemical parameters in cotton aphid. Results indicated that Sugar, glycogen, total protein and lipid in aphids have significant changes. Studies showed that among three strains called Ag-R, Ag-M and Ag-D which have different susceptibilities to neonicotinoids, strain Ag-R was the most tolerated aphids in counter of imidacloprid and thiametoxam, strain Ag-D was somewhat more tolerated to these insecticides and strain Ag-M was the most sensitive strain which has the resistance factor of nearly 890 in starvation method and was different depending on the method of exposure. Among energy sources, the total lipid in susceptible strain were decreasing more than resistant strain, whereas total proteins were increasing in resistant strain compared with sensitive strain. Total Glycogen was affected significantly in the stress condition which caused an increase and was the most resistant strain.     

河南不同地区棉蚜种群共生菌Wolbachia的检测及系统发育分析

【目的】本研究旨在揭示河南省不同地区棉蚜Aphis gossypii种群共生菌Wolbachia的感染情况,明确Wolbachia的感染类型及分类地位。【方法】2016-2017年间采集河南省不同地区的13个棉蚜种群,通过扩增COI基因片段对其进行种类鉴定;通过扩增棉蚜种群中Wolbachia的wsp基因片段对其进行Wolbachia感染率的检测,应用neighbor-joining法构建系统进化树进行棉蚜种群中Wolbachia的系统发育分析。【结果】对河南省内不同地区采集的13个棉蚜种群的Wolbachia感染率而言,郑州(ZZ)种群最高(46.67%),信阳2(XY2)种群最低(6.67%),13个种群Wolbachia的感染率范围为6.67%～46.67%,平均感染率为28.35%。基于wsp基因构建的系统发育树表明,安阳和信阳的棉蚜种群感染的Wolbachia株系属于B大组,其余地区棉蚜种群感染的Wolbachia株系属于A大组。【结论】河南省不同地区的棉蚜种群Wolbachia感染率差别较大,且不同种群感染的Wolbachia株系分别属于A大组或B大组。     

A contribution to Gymnosphaera (Cyatheaceae) in mainland Asia: Two new species,reinstatement of Cyathea bonii,and their phylogenetic positions

Gymnosphaera represents a minor lineage within the scaly tree-fern family Cyatheaceae. Tropical and subtropical mainland Asia is a main distribution area of Gymnosphaera. However, the species diversity of Gymnosphaera is currently incompletely known in mainland Asia due to lacking critical revision. Here we present new findings of species diversity and their relationships to mainland Asian Gymnosphaera based on field surveys, the examination of herbarium collections, and phylogenetic analyses of sequences of multiple chloroplast and nuclear regions. Two new species, G. saxicola from southwestern Yunnan and G. bachmaensis from central Vietnam, are established. Traditionally recognized G. podophylla is revealed to be a complex, from which G. bonii is reinstated as a distinct species. Our phylogenetic analyses identified four clades within Gymnosphaera in mainland Asia: G. denticulata clade, G. gigantea clade, G. podophylla clade, and G. salletii clade. The new species G. saxicola, which is special for its saxicolous habitat, was resolved as sister to G. austroyunnanensis in the G. salletii clade. The newly discovered G. bachmaensis, which is characterized specially by the spathulate frond, was positioned in the G. podophylla clade, being sister to G. bonii. The mountainous region from south-central Vietnam northwards to western Yunnan is a diverse center of Gymnosphaera and more species of this group are probably to be discovered there.     

Relationship between susceptibility to neonicotinoids and population dynamics of cotton aphid,Aphis gossypii Glover (Hemiptera: Aphididae)

Efficiency of imidacloprid and thiametoxam on population growth parameters of Aphis gossypii Glover (Hemiptera: Aphididae) was assessed using three bioassay methods including; residual, starvation and FAO dip methods. Aphids were assayed under laboratory conditions at 23 ± 2°C, photoperiod of 16:8 (L:D) and 70% relative humidity. Aphids were transferred to sprayed leaves and Petri dishes in residual and starvation methods, respectively. There was no feeding for the aphids in starvation method. In FAO dip method, insects were dipped for 5 s in pesticide solutions and then transferred to fresh leaves. Results revealed that LC₅₀ values calculated with the starvation and residual methods were respectively 15 and 11% more than FAO dip method (for imidacloprid) and 22 and 18% (for thiametoxam). The LC₅₀ value in starvation method was 3% more than the residual method. Calculated LC₅₀ in starvation and residual methods with imidacloprid respectively caused 160 and 34% increase in intrinsic rates of increase (r _m) and net reproductive rate (R ₀) in comparison with FAO dip method. Generation time (T) and doubling time (DT) were respectively 59 and 62% less than those in FAO dip method. In contrast, thiametoxam (LC₅₀ concentration) in starvation and residual method lead to 9 and 67% increase in r _m and R ₀ parameters compared to FAO dip method. However T and DT were respectively 65 and 92% less than mentioned parameters in FAO dip method. There was not any significant difference between life table parameters calculated by residual and starvation bioassay methods.     

Characterization of polymorphic microsatellite loci in the aphid species Macrosiphum euphorbiae (Hemiptera: Aphididae)

Fourteen microsatellite loci were isolated and characterized in the potato aphid, Macrosiphum euphorbiae Thomas, by screening a genomic library with the oligonucleotide probes (GA)₁₀ (GT)₁₀ and (GATA)₄. Allelic diversity was estimated in samples collected from potato fields in Tunisia. Ten loci displayed polymorphism that ranged from two to four alleles per locus and the observed heterozygosity ranged from zero to one. These markers could be used to study the population genetic structure of this polyphagous aphid species.     

Polymorphic microsatellites loci in the black Aphid,Aphis fabae Scopoli, 1763 (Hemiptera,Aphididae)

Aphis fabae is a widespread pest in Europe. This species includes four morphological cryptic subspecies which are mostly identified by their partially distinct secondary host range. To determine the extent of gene flow and isolation between these four taxa, we isolated and characterized eight microsatellite loci from A. fabae fabae and tested them on the three other subspecies. Polymorphism ranged from five to 35 alleles on the 138 individuals collected. When analysing each taxon separately, significant deviations from Hardy‐Weinberg (HW) equilibrum were observed. This could have different explanations, including the presence of null alleles, or the Wahlund effect or the mating system.     

Phylogeny of the genus Aphis Linnaeus, 1758 (Homoptera: Aphididae) inferred from mitochondrial DNA sequences

Aphis is the largest aphid genus in the world and contains several of the most injurious aphid pests. It is also the most reluctant aphid genus to any comprehensive taxonomic treatment: while most species are easily classified into "species groups" that form well defined entities, numerous species within these groups are difficult to tell apart morphologically and identification keys remain ambiguous and mostly rely on host plant affiliation. In this paper, we used partial sequences of COI/COII and CytB genes to reconstruct the first phylogeny of Aphis and discuss the present systematics. The monophyly of the subgenus Bursaphis and of the tree major species groups, Black aphid, Black backed aphid and frangulae-like species was recovered by all phylogenetic analyses. However our data suggested that the nominal subgenus was not monophyletic. Relationships between major species groups were often ambiguous but "Black" and "Black backed" species groups appeared as sister clades. The most striking result of this study was that our molecular data met the same limits as the morphological characters used in classifications: mitochondrial DNA did not allow the differentiation of species that are difficult to identify. Further, interspecies relationships within groups of species for which taxonomic treatment is difficult stayed unresolved. This suggests that species delineation in the genus Aphis is often ambiguous and that diversification might have been a rapid process.     

Suitability of Aphis craccivora (Hemiptera: Aphididae) and Bemisia tabaci (Hemiptera: Aleyrodidae) biotype-B as prey for Chrysopa pallens (Neuroptera: Chrysopidae)

The suitability of Aphis craccivora Koch (Hemiptera: Aphididae) and Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae) biotype-B eggs and nymphs as prey for pre-imaginal development and survival, adult longevity, and fecundity of the lacewing, Chrysopa pallens (Rambur) (Neuroptera: Chrysopidae) was evaluated under laboratory conditions at 25±1°C, 50±10% RH and a photoperiod of 16 h L:8 h D. Survival of C. pallens from first instar to adult eclosion was significantly different between the larvae that fed on the two prey species. C. pallens fed on A. craccivora completed development from egg to adult emergence, but those fed on eggs and nymphs of B. tabaci could not complete development, resulting in abnormal pupae and no normal adults emerged. The net reproductive rate (R ₀), intrinsic rate of natural population increase (r _m ), finite rate of increase (λ), mean generation time (T), index of population trend (I), doubling time (DT), and gross reproductive of rate (GRR) of C. pallens that fed on A. craccivora were 201.9 eggs per female, 0.13/d, 1.1/d, 40.1 d, 68.5, 5.2 d, 203.1 eggs per female, respectively. These results could be useful for mass-rearing of C. pallens and for understanding its population dynamics in the field in relation to the availability of different prey species.     

The distribution of aphids in response to variation among individual host plants: Uroleucon rudbeckiae (Homoptera: Aphididae) andRudbeckia laciniata (Asteraceae)

Abstract. 1. An investigation was conducted to determine the effects of variation among individual plants of Rudbeckia laciniata L. on the distribution and abundance of the aphid Uroleucon rudbeckiae (Fitch).
2. Wild plants were examined repeatedly for up to 2¹/₂ years to obtain information about plant growth and flowering, and about aphid densities on individual plants.
3. Aphid densities on individual plants were positively associated with several measures of plant vigour, including height, growth since previous year, and occurrence of flowers.
4. The colonization of plants by alate aphids was more highly correlated with plant height than with plant exposure. Aphid densities on individual plants were more strongly associated with plant height than with number of colonists.
5. In all years there were many hosts which were never colonized by aphids. Some of these plants produced shoots but did not flower. Experimental infestation of these plants showed that they were unsuitable for aphid growth and reproduction. Some plants which did flower and which escaped aphid infestation were nevertheless suitable hosts. Previously unsuitable host individuals were able to support aphids when grown under glass-house conditions.