Migration of alate morphs of the bird cherry aphid (Rhopalosiphum padi) and implications for the epidemiology of barley yellow dwarf virus

Suction trapping data indicate three periods of migration of Rhopalosiphum padi in spring, summer and autumn. Four alate morphs are present at different times during the year. A comparison of data from suction traps operating at 12·2 and 1·5 m suggests a different behaviour of females in autumn with more being recorded at 12·2 than 1·5 m. Males, which are only present in autumn, were also more numerous at 12·2 m. During tests to measure barley yellow dwarf virus (BYDV) infectivity, only 9% of female R. padi reproduced on oat seedlings in autumn compared with 74% in summer. Tests on alate female R. padi trapped alive showed that in summer all were exules, but during the first half of September these were largely replaced by gynoparae so that in autumn only 5% of all R. padi trapped at 12·2 m were alate exules. The aerial densities of gynoparae and males were 10 times greater at 12·2 than 1·5 m while densities of alate exules were similar at both heights. It is suggested that gynoparae and males fly higher to increase the chance of finding a taller dispersed host plant. The implications for BYDV epidemiology of the behaviour and presence of the various R. padi alate morphs indicate that autumn-sown cereals emerging before mid-September are particularly at risk from colonisation by alate exules before the transition to a mainly sexual migrant population is complete. Alate exules introduce BYDV from comparatively local sources. The ratio of total R. padi to Sitobion avenae in suction trap samples in autumn usually exceeds 100: 1, but on crops it was only 10: 1. The ratio of alate exule R. padi to S. avenae in suction traps in autumn was only 12: 1, similar to that observed on crops.     

Host plant selection by migrant Rhopalosiphum padi in autumn and the occurrence of an intermediate morph

Alate female Rhopalosiphum padi (L.) were trapped alive in suction traps in autumn, 1986–1988. Alate exules and gynoparae were identified by the morph of their offspring. Gynoparae largely replaced alate exules in September indicating that most of the population were holocyclic even in southern England where the primary host, Prunus padus L., is scarce. However, a few alate exules occurred throughout the autumn indicating that a proportion of the population is anholocyclic. A morph intermediate between alate exules and gynoparae occurred at low frequency throughout the autumn. Alate exules and gynoparae preferred barley and P. padus, respectively, on which to settle and larviposit; gynoparae were less selective than exules possibly due to an incomplete physiological switch from a preference for Gramineae to P. padus on which the sexual cycle is completed. Intermediate individuals preferred to settle on P. padus, but their virginoparous and oviparous nymphs were produced on barley and P. padus respectively. Intermediate morphs maintained under continuous short day conditions (L12 : D12) produced mainly apterous virginoparae with males occurring at the end of their reproductive live; intermediates occurred occasionally amongst the apterae. It is proposed that intermediates are a programmed component of the morph structure and have the potential to contribute to both the holocyclic and anholocyclic portions of the population.

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Résumé Des femelles de R. padi L. ont été capturées vivantes dans des pièges a succion en 1986–88. Les ailés exules et les gynopares ont été identifiés par le morphe de leurs descendants. Les gynopares ont remplacé généralement les ailés exules en septembre, ce qui montre que la majorité de la population est holocyclique même dans le sud de l'Angleterre où l'hôte primaire, Prunus padus L., est rare. Cependant, quelques ailés exules se maintiennent pendant l'automne montrant qu'une partie de la population est anholocyclique. Au cours de l'automne, on observe un morphe intermédiaire entre les ailés exules et les gynopares. Les exules ailés et les gynopares préfèrent respectivement l'orge et P. padus, sur lesquels ils s'installent et produisent des larves; les gynopares étaient moins sélectifs que les exules, peut-être par suite d'une rupture physiologique incomplète de leur préférence pour l'orge sur laquelle ils se développent rendant moins impérative leur attraction par P. padus sur lequel leur cycle sexuel est complet. Les individus intermédiaires préfèrent s'établir sur P. padus, mais leur larves virginipares et ovipares sont produites respectivement sur oreg et sur P. padus. Les morphes intermédiaires maintenus en jours courts (L12/N12) ont donné surtout des virginipares aptères les mâles apparaissant à la fin de leur vie; des intermédiaires sont apparus occasionnellement parmi les aptères. Les intermédiaires seraient une composante programmée de la structure polymorphe et contribueraient à la fois à l'apparition des fractions holocyclique et anholocyclique de la population.

     

Effects of the fungus <Emphasis Type="Italic">Lecanicillium lecanii</Emphasis> on survival and reproduction of the aphid <Emphasis Type="Italic">Schizaphis graminum</Emphasis>

This study investigated the effects of a strain of the fungus Lecanicillium lecanii (Zimm.) Zare and Gams (Hypocreales: Ascomycota) on the aphid Schizaphis graminum (Rondani) (Hemiptera: Aphididae). The fungus was administered to fourth instar nymphs and to alate and apterous adult morphs as a ground rice-kernel formulation. This study showed that L. lecanii formulation affected the survival of the aphids and interacted differently with the studied morphs, the Lethal Time values being lower for alate compared to apterous morphs and nymphs. The treatment also caused a significant reduction in the fecundity of the three treated aphid groups. Histological analysis revealed that the hyphae invaded the host hemocoel of a limited number of alate and apterous morphs: the fungus only entered through the spiracles. Scanning Electron Microscope observations revealed that L. lecanii adhered to the body surface of both adults and nymphs without differentiation. In conclusion, the present study suggests that this strain might be a good candidate for a programme of biological control of S. graminum and other aphid species.     

Morph determination in the bird cherry-oat aphid, Rhopalosiphum padi L

Both crowding and poor nutrition induce the appearance of emigrants in Rhopalosiphum padi L. No emigrants developed when the aphid was reared in isolation for six successive generations on actively growing leaves of bird cherry. However, emigrants developed as soon as the leaves ceased to grow or when the aphids became crowded. Crowding of mothers and postnatal crowding of the nymphs of apterous exules both influenced the induction of alate exules. The highest proportion of alate exules developed when both mother and offspring were crowded. Short day-length induced the appearance of gynoparae and males. For 50% production of gynoparae and males, over the range of temperature 10–18d?C, a 1.75 h reduction in day-length is required for every 4d? increase in temperature. At 18 d?C, even at day-lengths as short as 6 h, exposure for three generations is required before all offspring become gynoparae or males. Low temperature and short day-length was ineffective in inducing the development of gynoparae or males in the first three generations developing from the fundatrix.     

Performance of clones and morphs of two cereal aphids on wheat plants with high and low nitrogen content

The great variability of the aphid life cycle and their tendency for host alternation gives rise to aphid clones and morphs. Inter‐ and intraclonal variability may be observed in the responses of aphids to various environmental factors. In this study we aimed to evaluate the influence of intrinsic factors (clone and morph type) on the intrinsic rate of increase (r_m) of the English grain aphid, Sitobion avenae (Fabricius), and the bird cherry‐oat aphid, Rhopalosiphum padi (Linnaeus). For each species four apterous clones were collected from established laboratory colonies and compared to assess their relative fitness on high‐ and low‐nitrogen wheat plants under laboratory conditions. The clones had significantly different intrinsic rates of increase on high‐ and low‐nitrogen plants. All R. padi clones had a higher intrinsic rate of increase and mean relative growth rate than S. avenae. Experiments were also conducted to compare the mean fecundity of apterous and alate morphs of S. avenae and R. padi clones on high‐ and low‐nitrogen wheat plants. On high‐nitrogen plants the apterous morphs of S. avenae clones had significantly higher mean fecundity than alate morphs. There were no significant differences between the mean fecundity of alate morphs of the same species on high‐ and low‐nitrogen plants. The results support the idea of better fitness of specific clones/morphs on certain host plants due to higher and lower intrinsic rates of increase.     

Reproductive stragegies of the alate and apterous morphs of the grain aphid, Sitobion avenae

Differences were observed in the total fecundity and patterns of reproduction of four morphs of Sitobion avenae (F.). The reproductive rates of apterous and alate exules declined gradually with time but decreased rapidly in gynoparae and oviparae. Under short daylengths apterous exules gave birth in three phases; the first consisting mainly of gynoparae, the second of males and the third of non-gynoparous viviparae. The strategic importance of the sequence of morphs and the reproductive behaviour of each is discussed in relation to the maximization of overwintering population size.

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Stratégies reproductives des types ailés et aptères de Sitobion avenae F.

Résumé Des différences ont été observées entre les fécondités totales et les modes de reproduction de quatre types de Sitobion avenae. Les taux de reproduction diminuent graduellement dans le temps chez les aptères et ailés virginopares et rapidement chez les gynopares et ovipares. En photophases courtes, les virginopares aptères donnent naissance à trois types: le premier composé essentiellement de gynopares, le second de mâles et le troisième de vivipares. L'importance stratégique de la succession des types et du comportement reproducteur de chacun est discutée en relation avec l'accroissement maximum de la taille de la population hivernante.

     

The influence of clone and morph on the parameters of intrinsic rate of increase in the cereal aphids Sitobion avenae and Rhopalosiphum padi

This study aimed at evaluating the influence of intrinsic factors (clone, morph) on the parameters of the intrinsic rate of increase of Sitobion avenae (F.) and Rhopalosiphum padi (L.). For each species, apterous and alate exules of three clones originating from French oceanic regions were compared at 20 °C.The clonal factor had a significant effect on both time from birth until onset of reproduction (TBR) and intrinsic rate of increase (r). This effect was lower than that of the morph. Moreover, in the case of R. padi, a strong clone-morph interaction was observed.Both apterous and alate morphs were shown to have a higher rate of increase in R. padi than in S. avenae.Various models were fitted to aphid daily rate of reproduction. Both types of model and quality of fit depended mainly on the morph.

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Résumé Cette étude a eu pour but d'évaluer l'importance de facteurs intrinsèques (clone et morphe) sur les paramètres du taux d'accroissement de Sitobion avenae (F.) et Rhopalosiphum padi (L.).Pour chaque espèce, trois clones originaires de régions océaniques françaises ont été comparés à 20 °C pour les morphes exules ailés et exules aptères.L'effet du clone est significatif sur l'âge reproducteur (durée comprise entre la naissance du puceron et le début de sa phase de reproduction) et sur le taux intrinsèque d'accroissement naturel. Il est plus faible que celui du morphe avec lequel il intéragit dans le cas de R. padi.Les différences interspécifiques relevées mettent en évidence les capacités d'accroissement supérieures de R. padi par rapport à celles de S. avenae, dans le cas des ailés comme dans celui des aptères.Enfin, différents modèles ont été appliqués aux données de reproduction journalière des pucerons. Le type et la qualité des ajustements varient essentiellement en fonction du morphe.

     

Secondary host preferences and reproductive activity of the bird cherry-oat aphid, Rhopalosiphum padi

Of 15 species of grass, including cereals, alate exules and emigrants of Rhopalosiphum padi preferred to colonise Lolium perenne. Emigrants colonised cereals twice as readily as alate exules. Apterous exules also preferred L. perenne to cereals. Although small when reared on L. perenne, R. padi was more fecund per unit weight than on any of the other grasses tested. On the later growth stages of Dactylis glomerata and L. perenne, R. padi developed faster and were more fecund than on Phleum pratense at the same growth stages. At low temperatures R. padi were more fecund on P. pratense than on L. perenne and D. glomerata and vice versa at high temperatures. The secondary host preferences and reproductive activity of R. padi are discussed in relation to possible suitable hosts for viviparous overwintering and the pest status of this aphid in Britain.     

Detection of fungal metabolites of various Trichoderma species by the aphid Schizaphis graminum

The feeding preferences of alate and apterous morphs of the aphid Schizaphis graminum (Rondani) (Homoptera: Aphididae) were evaluated using leaves treated with powdered rice cultures of four fungal isolates belonging to different species of the genus Trichoderma (Deuteromycotina: Hyphomycetes). All of the fungal isolates restrained alate morphs of S. graminum from visiting treated leaves, but only Trichoderma citrinoviride Bisset also influenced the preference of apterous morphs. Trials carried out with supernatants obtained by centrifuging aqueous suspensions of the fungal cultures showed that the feeding preference of aphids was maintained in the absence of fungal spores and mycelia, supporting the hypothesis that at least part of the fungal metabolites responsible for this effect were water‐soluble compounds. Electrophysiological studies showed that the structures involved in the perception of the fungal metabolites are located on the aphid tarsomeres.     

The effect of temperature and wind on the flight activity of cereal aphids

The lower temperature threshold for take-off in Sitobion avenae obtained from an analysis of daily 12·2 m suction trap catches was 16°C. In the laboratory, the take-off threshold for S. avenae was 17·5°C in increasing temperatures, but 19°C when aphids were kept at constant temperatures for a short period of time. The equivalent thresholds were both 20·5°C for Metopolophium dirhodum. Over a period of 16 h no S. avenae took-off from plants at 10°C, 70% at 15°C and all within 16 h at 20°C. It was concluded that suction trap catches can be used to compare the temperature thresholds for take-off of both different species and morphs of a species of aphid. Emigrants of Rhopalosiphum padi, but not of M. dirhodum, took-off at a higher temperature than the alate exules. High winds were found to delay but not inhibit take-off both in the field, and in the laboratory, using both artificial and plant substrates. All aphids eventually flew, even from favourable hosts. It was concluded that adverse weather conditions will delay but not prevent cereal aphid migration in early summer.     

Description of oviparous females and alate males of Macrosiphum dzhibladzeae Barjadze, 2010 (Hemiptera: Aphididae) from Georgia

The hitherto unknown oviparous females and alate males of M. dzhibladzeae Barjadze, 2010, living on Euphorbia macroceras Fish. & Mey. and Euphorbia sp. (Euphorbiaceae), are described from resort Bakhmaro (Chokhatauri district, Guria region, Western Georgia). The life cycle of this species is established. Apterous and alate viviparous females and oviparous females of M. dzhibladzeae are compared to the same morphs of morphologically similar M. meixneri Börner, 1950. A key is provided to Euphorbia-feeding Macrosiphum spp. based on males.     

Body colour and genetic variation in winged morph production in the pea aphid

Aphids (Homoptera: Aphidoidea) produce a number of different phenotypes in their life-cycle, among which are winged (alate) and wingless (apterous) morphs. Lowe & Taylor (1964) and Sutherland (1969a, b) were the first to suggest that aphid clones differ in their propensity to produce the winged morph and that in the pea aphid (Acyrthosiphon pisum Harris), this propensity is linked to the colour of the phenotype. We tested for the occurrence of genetic variation in winged morph production by rearing individuals from red and green clones of pea aphid under wing-inducing (crowding) and control conditions, and scored the phenotypes of their offspring. Clones differed significantly in alate production and red clones produced on average a higher proportion of winged morphs than green clones. Importantly, however, there was considerable variation between clones of the same colour. Broad-sense heritabilities of winged morph production were 0.69 (crowding treatment) and 0.63 (control). Clones also differed in the number of offspring they produced. When exposed to the crowding stimulus, aphids deferred offspring production, resulting in a higher number of offspring produced in the crowding treatment than in the control.     

Water loss in desert ants: caste variation and the effect of cuticle abrasion

Water loss rates (WLRs) varied across castes (workers > alate males > alate females) for desert ants in the genera Aphaenogaster, Messor and Pogonomyrmex. Exposure to soil caused increased WLRs in workers and foundresses (mated, dealate females), apparently because of cuticular abrasion caused by nest excavation. Moreover, field‐collected workers and foundresses from incipient nests (those exposed to soil) had similar WLRs, as did unabraded workers and alate females (those unexposed to soil). These data call into question previous adaptive scenarios for differences in WLRs across ant species and castes. For live alate females, WLRs increased over two stages. The first increase occurred immediately after mating, and the second occurred for foundresses collected 2 days later from incipient nests. By contrast, WLRs of HCN‐killed females were unaffected by mating, but increased significantly for foundresses collected from 2‐day‐old nests.     

Phenology of the black bean aphid,Aphis fabae,on organic crops and effect of parasitoid communities

1. Parasitoids represent the third trophic level in plant-insect food webs. They develop in or on herbivorous insects at the second trophic level, which in turn feed on plants that constitute the first trophic level. This food web system might be affected by agricultural landscape and practices.
2. We studied the phenology and population density of Aphis fabae (Hemiptera: Aphididae) and determined the impact of parasitoids in organic fields of beans and beet plants.
3. Twenty leaves per field were collected weekly. For each leaf, the total number of aphids, apterous and alate morphs, the number of mummies, and mortality of aphids were recorded.
4. Our results showed that the phenology of A. fabae varied throughout the season with maximal numbers observed at the beginning of summer. In addition, the production of alate morphs and the proportion of parasitism were positively correlated to an increase in aphid densities.
5. Six parasitoid species were recorded attacking A. fabae in the field, with Lysiphlebus fabarum and Binodoxys angelicae (Hymenoptera: Aphidiinae) being the most abundant parasitoids.
6. Field characteristics and availability of host plants in the surroundings affected A. fabae density. Lysiphlebus fabarum and B. angelicae appeared to be good biocontrol candidates against A. fabae.

     

The effect of starvation and subsequent feeding on the reproductive potential of the grain aphid,Sitobion avenae

The short‐term starvation tolerance of alate and apterous morphs and the effect of periods of starvation on the longevity and fecundity of alate adults were evaluated in the grain aphid, Sitobion avenae (Fabricius) (Hemiptera: Aphididae). Alate adults exhibited a proportionally larger length of survival compared with apterous adults under continuous starvation conditions. Newly molted pre‐reproductive adults were starved for 0–96 h and their survival rate on the 1st day after recovering with food was not significantly different from that of control aphids. Starvation reduced lifetime fecundity, but increased the reproductive rate immediately after nutrition being improved. Fecundity and longevity after 24, 48, 72, and 96 h of starvation were significantly higher than after 120 or 144 h of starvation. However, no significant differences were observed for alate adults after 24, 48, 72, or 96 h of starvation. This study suggests that the ability of alatae to adapt to brief periods of starvation could be one of the important factors affecting the reproductive success of aphids during delays in locating host plants.     

Geographic variation in Pemphigus populitransversus (Insecta: Aphididae)*†

The geographic variation of 33 morphological characters of two morphs of the gall-forming aphid Pemphigus populitransversus is studied in 214 locality samples. Among-locality variation ranges from 1 to 69% in the elongate morph and from 0 to 44% in the globular morph. The design of the study permits separation of interlocality correlations from intralocality correlations. The former are partly a function of the latter, confirming early observations on another Pemphigus species and on ticks. Factor analyses of both correlation matrices for both morphs yield four factors. Within localities these factors agree for the two morphs; among localities only one factor corresponds. Multiple discriminant analyses among localities of the two data sets do not correspond. There is little correlation between characters of stem mother and alate morphotypes within localities but such correlations are strong among localities. Maps are furnished for characters representing the independent dimensions of variation for each morph. Patterns of variation are shown to be significant by spatial autocorrelation analysis for both morphs but are much more marked in the elongate morph. Significant positive autocorrelation occurs up to 1000 km in the elongate morph – mostly only up to 200 km in the globular morph. There are two to three geographic variation patterns in the elongate morph, whereas in the globular morph the classes of patterns are less well defined and involve few characters. The environmental factors to which the globular and elongate morphs are adapting would appear to have different autocorrelation patterns. In each morph the patterns are clearly different and cannot be explained by a single microevolutionary process. The findings are compared with an earlier study in the related and largely sympatric P. populicaulis.