Coexistence of three species of leaf-feeding aphids (Homoptera) onBetula pendula

Summary Three sympatric aphid species of European origin (Euceraphis betulae (Koch),Callipterinella calliptera (Hartig) andBetulaphis brevipilosa Börner) feed onBetula pendula Roth in northern California. The component species of this guild partition resources by plant part, feeding sites within leaves and leaf phenological state. Overall niche overlap between species reached a maximum of 49.9% betweenB. brevipilosa andC. calliptera. Co-occurence ofE. betulae with both other species on leaves was random while co-occurrence ofC. calliptera andB. brevipilosa was aggregated. Morphological features (stylet length and body weight) act to mediate niche organization within this guild.California populations ofE. betulae andB. brevipilosa appear to utilize resources in a fashion similar to endemic Danish populations. Whereas DanishC. calliptera usually feed inside leaves silked together by spiders or Lepidoptera,C. calliptera in California also commonly feed on exposed leaf surfaces. This species, which has thus expanded its endemic niche, occupies the most generalized niche of the three species studied.     

Effect of parasitism on flight behavior of the soybean aphid, Aphis glycines

Many aphid species possess wingless (apterous) and winged (alate) stages, both of which can harbor parasitoids at various developmental stages. Alates can either be parasitized directly or can bear parasitoids eggs or larvae resulting from prior parasitism of alatoid nymphs. Winged aphids bearing parasitoid eggs or young larvae eventually still engage in long-distance flights, thereby facilitating parasitoid dispersal. This may have a number of important implications for biological control of aphids by parasitoids. In this study, we determined the effect of parasitism by Aphelinus varipes (Hymenoptera: Aphelinidae) on wing development and flight of the soybean aphid, Aphis glycines (Hemiptera: Aphididae). We also quantified the influence of aphid flight distance on subsequent A. varipes development. Parasitism by A. varipes was allowed at different A. glycines developmental stages (i.e., alatoid 3rd and 4th-instar nymphs, alates) and subsequent aphid flight was measured using a computer-monitored flight mill. Only 35% of aphids parasitized as L3 alatoid nymphs produced normal winged adults compared to 100% of L4 alatoids. Flight performance of aphids parasitized as 4th-instar alatoid nymphs 24 or 48 h prior to testing was similar to that of un-parasitized alates of identical age, but declined sharply for alates that had been parasitized as 4th-instar alatoid nymphs 72 and 96 h prior to testing. Flight performance of aphids parasitized as alate adults for 24 h was not significantly different from un-parasitized alates of comparable ages. Flight distance did not affect parasitoid larval or pupal development times, or the percent mummification of parasitized aphids. Our results have implications for natural biological control of A. glycines in Asia and classical biological control of the soybean aphid in North America.     

Host selection, development, survival, and reproduction of turnip aphid (Homoptera: Aphididae) on green and red cabbage varieties

Host selection, development, survival, and reproduction of the turnip aphid, Lipaphis erysimi (Kaltenbach), on selected green and red cabbage varieties, Brassicae oleracea L. variety capitata, were studied in the laboratory. Although numbers of alate L. erysimi on different varieties varied generally, more alate adults and nymphs were observed on green than on red varieties on day 3; whereas, these differences were not so significant on day 15. There were no significant differences in alate adults on all six varieties on day 15. Numbers of nymphs and apterous adults of L. erysimi varied among the varieties. When mean numbers of those aphids on green varieties were compared with those on red varieties, the green varieties had more nymphs and apterous adults than those on the red varieties. Developmental durations of L. erysimi nymphs also varied among all varieties, and the nymphs fed on green varieties developed longer than those fed on red varieties. Adult longevities on these cabbage varieties were not significantly different; whereas, the adult fecundities were. Survival rates of L. erysimi varied greatly among all varieties, but there are no significant differences between the green and red varieties. Factors that might affect aphid host selection, growth, longevity, fecundity, and survivorship were discussed.     

Prey Preference of Aphidoletes Aphidimyza on Acyrthosiphon Pisum: Effect of Prey Color and Size

Color polymorphism in insects as well as factors contributing to its occurrence and maintenance have been of a great interest. Pea aphid (A. pisum) exhibits a noteworthy color polymorphism which occurs as red and green. The preference of the predatory gall midge Aphidoletes aphidimyza (Rondani) for the two color morphs of pea aphid at two life stages (adult and second instar) was investigated. Red adults, red nymphs, green adults, and green nymphs were offered to the larvae of the predator in different sets. The larvae attacked red aphids more than green ones of a same size. But whenever adults were offered along with nymphs, the nymphs experienced significantly more attacks. Although visual cues could result in more predation on red aphids, our findings showed that the size of aphids was the more important factor. The results showed that previous findings, suggesting more predation on the red morph, are valid when the same size of aphids is used. The ovipositing females exhibited no differences in oviposition choice between stalks infested with either of two colors.     

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.     

Effects of bacterial secondary symbionts on host plant use in pea aphids

Aphids possess several facultative bacterial symbionts that have important effects on their hosts'' biology. These have been most closely studied in the pea aphid (Acyrthosiphon pisum), a species that feeds on multiple host plants. Whether secondary symbionts influence host plant utilization is unclear. We report the fitness consequences of introducing different strains of the symbiont Hamiltonella defensa into three aphid clones collected on Lathyrus pratensis that naturally lack symbionts, and of removing symbionts from 20 natural aphid–bacterial associations. Infection decreased fitness on Lathyrus but not on Vicia faba, a plant on which most pea aphids readily feed. This may explain the unusually low prevalence of symbionts in aphids collected on Lathyrus. There was no effect of presence of symbiont on performance of the aphids on the host plants of the clones from which the H. defensa strains were isolated. Removing the symbiont from natural aphid–bacterial associations led to an average approximate 20 per cent reduction in fecundity, both on the natural host plant and on V. faba, suggesting general rather than plant-species-specific effects of the symbiont. Throughout, we find significant genetic variation among aphid clones. The results provide no evidence that secondary symbionts have a major direct role in facilitating aphid utilization of particular host plant species.     

Behavioral interactions between three birch aphid species and Adalia bipunctata larvae

First and fourth instars of three birch aphid species were exposed to first and fourth instars of Adalia bipunctata (L.), the most common aphid predator on silver birch, Betula pendula Roth, in northern California. Defensive behavior differed by aphid species. Euceraphis betulae (Koch) (Eb), the most successful escapee, was highly mobile and frequently walked away from coccinellid larvae. Betulaphis brevipilosa Börner (Bb), a flat, sessile species, was the least successful aphid at actively escaping from A. bipunctata larvae, but could passively escape detection when coccinellid larvae walked over nymphs and did not perceive them. Active escape behavior was much safer for aphids than passive avoidance of detection. Both instars of Eb and fourth instars of Callipterinella calliptera (Hartig) (Cc) escaped from coccinellid larvae more frequently when approached from the front, apparently using vision for pre-contact detection of A. bipunctata. These aphids avoided physical contact with larger predators more often than with smaller predators. Level of predation by A. bipunctata on these three aphid species is dependent upon types of aphid defense.

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Résumé Des larves de premier et troisième stades de 3 espèces de pucerons du bouleau: Betulaphis brevipilosa, Callipterinella calliptera et Euceraphis betulae ont été exposées aux attaques des larves du premier et du quatrième stades d'Adalia bipunctata, prédateur le plus fréquent de pucerons sur Betula pendula en Californie du Nord. Le comportement défensif dépend beaucoup de l'espèce de puceron. E. betulae, qui s'échappe avec le plus de succès, est très mobile et s'écarte fréquemment des larves agressives de coccinelles. B. brevipilosa, espèce plate, sessile, a le moins de succès dans la protection active contre les attaques des larves de A. bipunctata, mais elle peut échapper passivement à la détection des larves de coccinelles qui ne peuvent pas les discerner lorsqu'elles circulent parmi les larves de pucerons. E. betulae (aux deux stades) et C. calliptera (au quatrième) échappent le plus aux larves de coccinelles qui attaquent de front, ce qui laisse supposer que la vision permet de détecter A. bipunctata avant le contact. Le succès des larves de coccinelles varie selon les stades du puceron et de la coccinelle. Les possibilités d'obtention de niveaux de prédation élevés de ces 3 pucerons par A. bipunctata dépend du type de protection du puceron.

     

Diurnal pattern of aphid feeding and its effect on cotton leaf physiology

Cotton aphid (Aphis gossypii G.) populations seemed to fluctuate over the past years in cotton (Gossypium hirsutum L.) perhaps as a result of excessive use of insecticides for controlling more problematic pests. Contradictory plant responses have been observed depending upon the aphid/plant system, and it is unclear if cotton aphids, abiotic stress or both are responsible for cotton yield reduction in aphid-infested fields. Our objectives were to investigate the diurnal changes in the physiology of cotton leaves following aphid herbivory, and the diurnal pattern of aphid feeding. The experiment was conducted in a growth chamber using the cotton cultivar ‘Stoneville 474’. Leaves of the same age and size were infested with wingless adults plus nymphs. Cotton aphids were allowed to increase in numbers without restriction for 9 days, after which the amounts of carbohydrates in aphid-honeydew, and the number of honeydew droplets excreted per aphid were measured. Photosynthetic rates, dark respiration rates and foliar non-structural carbohydrates were measured. The amount of individual carbohydrates found in the honeydew was significantly different with time. The total amount of carbohydrates excreted per aphid within a 24-h period averaged 2.5 μg. The number of honeydew droplets excreted per aphid varied significantly from time to time period. Cotton aphids did not significantly alter photosynthesis or respiration rates or non-structural carbohydrates on leaves. Aphid populations of approximately 300 per leaf on the 9th day of infestation did not appear to significantly alter the physiology of cotton leaves.     

Aphids do not attend to leaf colour as visual signal, but to the handicap of reproductive investment

The evolution of visual warning signals is well known in animals but has received scant attention in plants. The coevolutionary hypothesis is the most influential hypothesis on warning signals in plants proposing that red and yellow leaf colours in autumn signal defensive strength to herbivores. So far, evidence in support of the hypothesis, which assumes a coevolutionary origin of autumnal leaf colours, is correlative and open to alternative explanations. We therefore tested the coevolutionary hypothesis experimentally by colouring the leaves either red or green of same-aged mountain ash (Sorbus aucuparia) individuals. We monitored the response of winged aphids to leaf colour using insect glue on branches with natural and artificial leaf colours in each individual. In contrast to the prediction of the coevolutionary hypothesis, aphid numbers did not differ between the individuals with artificial green or artificial red leaves. Likewise, at the within-plant level, aphids did not colonize branches with natural green leaves preferentially. However, we suggest that plants emitted warning signals because aphids colonized the hosts non-randomly. We found a strong positive correlation between aphid numbers and fruit production, suggesting an allocation trade-off between investment in plant defence and reproduction. Our study demonstrates that aphids use warning signals or cues in host selection, probably volatiles, but that they did not use leaf colour.     

Response of alate aphids to green targets on coloured backgrounds

To study the effect of background colour on aphid landing on green targets (water pan traps), two field experiments were set up in Hessen, Germany, in 2003. Traps were put onto coloured plastic sheets (13 colours, straw mulch, transparent foil, and uncovered soil, Experiment 1). In Experiment 2, green water pans were again put on coloured plastic sheets (red, white, green, and yellow), and the sheets were either sprayed with insect glue or not. Backgrounds and traps were spectrally characterised with a field radiometer (320–950 nm). Aphid catches were highest in the traps on the uncovered background, and lowest in the traps on white or silver backgrounds. For Brevicoryne brassicae, Myzus persicae (Homoptera: Aphididae, Macrosiphini) and five further aphid species, there was a significant negative correlation between UV‐reflectance (320–400 nm) and log(N + 1)‐transformed number of individuals. However, the effect of straw mulch (reduced aphid catches with straw compared to the uncovered background), could not be attributed to differences in UV‐reflectance, as UV was almost identical in soil and straw. High numbers of alate aphids were caught in traps with dark backgrounds (e.g. black, dark green), a result which was attributed to the high contrast between the background and target. The substantially higher aphid numbers from targets with bare soil than from targets with spectrally similar black backgrounds were thought to be caused by the structure of the background surface: for alate aphids, landing close to the target on smooth surfaces may induce probing, and the lack of appropriate substrate will result in take‐off, whereas soil will not induce probing, and aphids will continue to move towards the green targets.     

Influence of sticky trap color and height above ground on capture of alate Elatobium abietinum (Hemiptera: Aphididae) in Sitka spruce plantations

A series of field trials were used to assess the practicality of using sticky traps to monitor populations of green spruce aphid, Elatobium abietinum (Walker), in plantations of Sitka spruce. The highest numbers of alate E. abietinum were caught on sticky traps placed in the upper third of the live canopy at 9-17 m above the ground, whereas low numbers of aphids were caught just below the live canopy or at 2 m above the ground. Trials in 2005 with sticky traps of different colors showed that significantly more alate E. abietinum were caught on yellow, red, and green sticky traps than on white, blue, and black traps. A repeat trial in 2007 resulted in significantly more alate aphids being caught on red sticky traps than on traps of any other color except for green. Attraction to red is unusual among aphids, as aphids are thought not to possess a red-sensitive photoreceptor. The attraction of E. abietinum to red-colored sticky traps suggests that conifer-feeding aphids might have a fundamentally different color response compared with aphids that live on cereals, grasses, or herbaceous plants. Alternatively, the attraction to red might be a physiological artifact related to the presence of red-screening pigments in the aphid's compound eye.     

A controlled environment study of the effect of leaf physiological age on the movement of apterous Myzus persicae on sugar-beet plants

Apterous Myzus persicae were found to move frequently from leaf to leaf on sugar-beet plants in controlled environment conditions. It is suggested that aphid movement can be related to changes in the rate and content of translocate flow during leaf development. These changes make newly-emerged leaves nutritionally favourable to colonising aphids and make expanding leaves slowly wane in favourability during the process of ‘sink to source’ conversion leading to aphid dispersal from the leaf. Variation in temperature was not found to alter the rate of aphid movement or the period (measured in thermal time) that aphids spent on particular leaves. However, the lower temperature was found to increase the rate of aphid development, aphid size and fecundity; these effects could also be due to nutritional factors. This dispersal behaviour may be a tactic to maximise food intake by a polyphagous aphid and increase the probability that nymphs are deposited on nutritionally-favourable leaves. The implications of the interleaf dispersal of apterous M. persicae for within- and between-plant spread of beet yellows virus (BYV) and beet mild yellowing virus (BMYV) are discussed.     

Prevalence of Pandora neoaphidis (Zygomycetes: Entomophthorales) infecting Nasonovia ribisnigri (Hemiptera: Aphididae) on lettuce crops in Argentina

Lettuce crops, Lactuca sativa, organically produced in La Plata, Argentina, were sampled in order to determine the prevalence of fungal diseased aphids. Nasonovia ribisnigri was the only aphid detected and its occurrence was highly variable. The fungal pathogen Pandora neoaphidis (Entomophthoromycotina: Entomophthorales) was the only pathogen detected. We recorded a maximum of 34.2 aphids per plant and the highest rate of fungal prevalence was 56.6% (n = 30) (aphids infected/total aphids). Infected aphids were observed in all sampling sites. No differences of infection rates were detected between the center and the edge of crops. Host density was an important factor determining infection. The majority of host population was comprised of nymphs which were the most infected in terms of individuals per habitat unit (lettuce plant), but considering the proportion of infected aphids per stage of development, the prevalence of infection in nymphs and adults was similar.     

Electroantennogram responses of aphid nymphs to plant volatiles

Abstract. Electroantennogram (EAG) responses of immature aphids were investigated for 30 plant volatile compounds in third‐ and fourth‐stadium nymphs of the black bean aphid, Aphis fabae. The nymphs were destined to develop into adult alate (winged) virginoparae. The EAG response profiles were similar to those previously reported in the adults. Among the compounds tested, hexanonitrile elicited the largest EAG responses in both nymphal stadia, corresponding to previously reported results with adults. Six‐carbon aliphatic compounds showed relatively higher EAG activities in the nymphs but, in contrast, (E)‐2‐hexenal, benzaldehyde, α‐pinene and β‐pinene, and citronellal elicited relatively smaller EAG responses in nymphs than adults. Although overall EAG response profiles were similar between the third and the fourth stadia for the majority of the volatiles, four aldehyde compounds, hexanal (E)‐2‐heptenal, 2‐hydroxybenzaldehyde and citronellal, showed relatively higher EAG activities in the third than in the fourth stadium. The present study indicates that aphid nymphs possess a functional olfactory receptor system before the antennae are fully developed morphologically and physiologically.     

Interannual dynamics of aerial and arboreal green spruce aphid populations

Partial defoliation of spruce by the green spruce aphid Elatobium abietinum (Walker) is a recurrent event in European and, increasingly, North American forests. The patterns of insect abundance on trees have never been satisfactorily described by a numerical model despite considerable knowledge of endogenous and exogenous factors in the population dynamics of the species. Long-term field population estimates of the aphid on foliage provided the opportunity to evaluate such a model. Unlike comparable models for tree-dwelling aphids, this was also applicable to almost completely independent aphid field data derived from the Rothamsted Insect Survey’s nationwide network of suction traps. Although based on relatively few parameters, the model was robust in its predictions of alate aphids geographically remote from the forest in which the original population was estimated. The population maximum, which causes the greatest forest damage, is reached in early summer and can be predicted from knowledge of winter temperature (chill bouts), spring temperature (thermal sum), and interannual negative feedback (density dependence). The model provides confirmation that alate populations of spruce aphids, upon which a number of other extensive studies have been based, are ultimately influenced by similar endogenous and climatic factors and that they are a reasonable proxy for aphids on trees.     

Comparative transmission of pea mosaic virus by one strain ofMyzus persicae Sulz. and two strains ofAcyrthosiphon pisum Harris

Experiments were conducted to compare the efficiency of transmission of a strain of pea mosaic virus (PMV) isolated in Czechoslovakia by two strains (clones) of the pea aphidAcyrthosiphon pisum Harris (green and red) and one strain ofMyzus persicae Sulz. PMV is a nonpersistent virus and the preliminary fasting of aphids before acquisition feeding increases the efficiency of aphids in transmission of this virus. In our experiments two hour fasted individuals were used and two periods of acquisition feeding on the source (1 and 5 minutes). On the healthy test plants the aphids were left over night. As the source and test plants pea (Pisum sativum L.) of the cv. Raman were used. During the one minute acquisition period on the source of infection the aphids were observed under the stereoscopic microscope. They usually made two to three probes. During the five minute acquisition feeding time the aphids were not observed and they were taken from the source of infection after a lapse of five minutes. To compare the efficiency in transmission of this virus by these aphids only one aphid per tested plant was used and in all trials only two to four day old nymphs were taken. Differences in transmission efficiency between two strains ofAcyrthosiphon pisum Harris were highly significant. The green strain of pea aphid was the less efficient vector in comparison with the green peach aphid and the red strain of the pea aphid, the latter being the most efficient vector of this virus.     

桃蚜对不同单色光趋性反应的测定

为了探讨蚜虫对不同色光选择反应的定量指标, 采用滤光片技术测定了有翅和无翅桃蚜Myzus persicae对不同波长单色光的趋性反应。结果表明: 有翅蚜对490~550 nm范围内的单色光表现出明显趋性, 其中对538.9和549.9 nm的绿偏黄色光趋性最强, 平均位移分别达25.29和22.97 cm, 其次为491.5 nm的蓝绿色光, 而对于波长576.0 nm的黄色光并没有表现出明显趋性。无翅蚜对不同单色光的趋性反应则没有明显的峰值, 最高相对平均位移仅1.41。行为测定结果与前人电生理测定的结果基本一致, 说明以位移作为小体昆虫趋光性强弱的指标是可靠的。     

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.     

Do aphids paint the tree red (or yellow)—can herbivore resistance or photoprotection explain colourful leaves in autumn?

We explored two mutually nonexclusive hypotheses on autumnal leaf colouration. The co-evolutionary hypothesis states that autumnal leaf colouration functions as a handicap signal to herbivorous insects, whereas the photoprotection hypothesis posits that plant pigments primarily protect the plant against cold-induced photoinhibition and enhance nutrient transfer. To contrast both hypotheses, we compared yellow and red leaf colouration in three groups of mountain ash (Sorbus aucuparia L.). Two montane groups of different age were characterised by low aphid numbers and low temperature, and a lowland group by high aphid numbers and high temperature. There were no consistent altitudinal differences in leaf colouration. Compared to young trees, adult trees developed fewer red but more yellow leaves at high altitude. In the lowland population, the development of red leaf colour was related to decreasing daytime temperature, whereas the appearance of yellow leaf colouration corresponded to the decreasing photoperiod. This is consistent with the photoprotection hypothesis. Individual differences in red and yellow leaf colouration were inversely correlated to the number of fruits, which might be interpreted as a trade-off between reproductive and protective commitment. Temperature effects explained variation in aphid numbers over time and leaf colouration explained aphid distribution on a given day. As predicted by the co-evolutionary hypothesis, strongly coloured individuals harboured fewer aphids than green or dull-coloured ones. Since decreasing temperature reduced the number of migrating aphids but induced red leaf colouration, these processes are not mutually fine-tuned, which likely restricts the potential for co-evolution between mountain ash and aphids.     

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.