Change in response of Rhopalosiphum padi spring migrants to the repellent winter host component methyl salicylate

Olfactometry showed that the response of spring migrants of the bird cherry-oat aphid, Rhopalosiphum padi (L.) (Homoptera: Aphididae), to the repellent winter host volatile methyl salicylate changes with age of the adult aphid. Between three and four days after becoming adult, and having left the winter host Prunus padus L., aphids lost their negative response to the chemical. The change in response was not associated with contact with a summer host, oats. In a settling choice bioassay, migrants avoided oats which had been exposed to volatile methyl salicylate. Aphids with removed antennal tips did not avoid the exposed plant, indicating that plant choice was influenced by cues from the plant surface. The results are discussed in relation to the use of methyl salicylate in integrated control.     

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.

     

Odour stimuli affecting autumn migration of Rhopalosiphum padi (L.) (Hemipter: ornoptera)

R. padi gynoparae settling in the canopy of the winter host, Prunus padus, tended to form aggregations. Differences in the degree of exposure to wind, sun, etc., did not satisfactorily explain the aggregated pattern observed. In laboratory experiments, aphids did not distinguish between leaves taken from nearby aggregations and leaves taken far away from aggregations. The results of laboratory olfactometer tests and field trapping studies indicate that R. padi gynoparae produce an aggregation pheromone. Thus the aggregated settling pattern of gynoparae of R. padi can best be explained as a response to an aggregation pheromone.     

Are primary woody hosts ‘island refuges’ for host‐alternating aphids and important for colonization of local cereals?

Only few studies are available dealing with the relation between winter host density and spatial distribution and spring colonization of winter cereals by the host‐alternating cereal aphid species Rhopalosiphum padi and Metopolophium dirhodum. Large‐scale studies in climatically different agroecosystems in Germany from 2004 to 2006 revealed for R. padi and M. dirhodum larger spring/summer populations in landscapes with higher densities of winter hosts. A small‐scale study was performed in winter wheat fields adjacent to a large hedge with several typical winter hosts plants, bird cherry (Prunus padus) and wild rose species (Rosa spp.) to indentify distance effects (0–8, 8–24 and 24–60 m). Weekly measurements of aphid density between May to July showed significantly higher densities of R. padi compared with those of other aphids. Statistical analysis (Tukey–Kramer test and regression analyses) revealed significant gradients from the hedge to the field centre for R. padi and M. dirhodum. In comparative studies, winged R. padi from winter and adjacent summer hosts were genotyped using four microsatellite markers. The results showed that individuals from a certain winter host were not genetically similar with individuals from neighbouring summer hosts; it, therefore, seems that winter host clones did not significantly contribute to population built‐up in cereal fields over short distances. It could be concluded that on a regional scale, the density of sources for early migrants of R. padi is important for colonization intensity of surrounding summer hosts, but that the high local movement intensity and the relative small proportion of aphids that could be analysed in such tracking studies are blurring close spatial relations within short time periods.     

Host monitoring by aphid migrants: do gynoparae maximise offspring fitness?

Summary Gynoparae (autumn migrants) of the bird cherry-oat aphid,Rhopalosiphum padi (L.), were not found to land randomly on their primary host,Prunus padus L. Some trees, although within a few metres of heavily infested trees, were not colonized at all. This phenomenon occurs regularly from year to year.The reasons for this non-random landing pattern are examined in terms of the success of subsequent generations, special attention being paid to the reproductive rate of the oviparae and fundatrices and the success of the developing populations in the spring of the following year.Trees in Scotland and Finland were examined in this manner and the hypothesis developed that the gynoparae ofR. padi show maternal care in selecting hosts that favour their offspring's survival and reproduction.Aphids were reared under controlled conditions in the laboratory on twigs cut from trees of known previous infestation levels. Aphids on twigs from preferred trees had better survival rates and produced more offspring than those aphids on twigs from non-preferred trees.     

Larval food affects oviposition preference,female fecundity and offspring survival in Yponomeuta evonymellus

1. Yponomeuta evonymellus is a monophagous moth that feeds on Prunus padus which is native to Europe. In recent years, larval feeding and egg clusters have also been observed on non‐native Prunus serotina plants; however, survival of larvae on this new host is very low. 2. The objective of the present study was to determine how the feeding of larvae on each of the two host plants impacts oviposition, offspring survival and fecundity in Y. evonymellus. Our hypothesis was that, under controlled conditions, females will lay eggs on the host on which they fed as larvae. We also hypothesised that the lower survival of young larvae feeding on P. serotina was due to the smaller buds and leaves present in this species, relative to those of P. padus. 3. A dual‐choice experiment conducted under laboratory conditions demonstrated that females preferentially chose to oviposit on the plant species on which they fed as larvae. In the experiment, potential fecundity and offspring survival were significantly higher on P. padus than on P. serotina. The reduced performance of Y. evonymellus on P. serotina was correlated with a smaller bud mass and volume, lower leaf mass and surface area, and difficulty in constructing a protective tent against unfavourable weather conditions. 4. In summary, the identity of the host plant species during larval feeding determines adult oviposition preference for that host species. The survival of larvae on P. serotina growing in the nature is low, but for phenology‐related reasons.     

Chemical aspects of host‐acceptance behaviour in the bird cherry‐oat aphid Rhopalosiphum padi: host‐acceptance signals used by different morphs with the same genotype

Host acceptance by gynoparae and winged virginoparae of the bird cherry‐oat aphid Rhopalosiphum padi (L.) is investigated utilizing leaves and aqueous extracts of the primary and secondary hosts, as well as nonhost plants. Gynoparae are specialized to reproduce on bird cherry Prunus padus L., whereas virginoparae reproduce and feed on various grasses. Host acceptance is assessed using levels of reproduction and survival for adults, as well as survival for nymphs. Little is known of host acceptance by nymphs. The data show that gynoparae and winged virginoparae produce nymphs almost exclusively on their host plants, bird cherry and barley leaves, respectively, over 72 h. When tested with aqueous plant extracts, however, gynoparae produce nymphs almost exclusively on bird cherry extract and progeny numbers are found to be similar to those on plant leaves. Few nymphs are produced on artificial diet. By contrast, winged virginoparae produce nymphs on aqueous extracts of barley, bird cherry and bean, as well as on artificial diet. The numbers of nymphs deposited by gynoparae are similar on aqueous extracts of bird cherry leaves collected at different times during the growing season. When extracts from leaves of various Prunus species are tested, only leaves of P. padus and Prunus virginiana stimulate parturition. Oviparae, the sexual female nymphs of gynoparae, survive well for 96 h on both bird cherry and barley leaves but not on bean seedlings, whereas nymphs of winged virginoparae survive well only on barley leaves. They do not survive for 96 h on any plant‐leaf extracts, although they do survive on artificial diet.     

Food acceptance by a monophagous and an oligophagous insect in relation to seasonal changes in host plant suitability

Food acceptance by larvae of two lepidopteran species feeding on Rosaceae, viz. Yponomeuta evonymellus (monophagous) and Y. padellus (oligophagous), was compared. The influence of seasonal changes in plants as food for both insects was examined, in particular, the effects of nitrogen and sorbitol in leaves. In the laboratory, Y. evonymellus accepts Crataegus monogyna, a host plant of Y. padellus, and Y. padellus accepts Prunus padus, the host plant of Y. evonymellus. P. padus is the most suitable food plant for Y. evonymellus. No difference in food-quality for Y. padellus was found between C. monogyna and P. padus. The performance of both species on P. padus is less influenced by seasonal changes than on Crataegus. The suitability of Crataegus decreases during the season. This is probably caused by the decrease of its nitrogen content, and not by the decrease of sorbitol in the plant. The monophagous, Y. evonymellus, is more sensitive to seasonal changes in its food when fed with a non host plant than the oligophagous Y. padellus. In oviposition experiments both species have a preference for their normal host-plants.     

Adaptation of a generalist moth, Operophtera brumata, to variable budburst phenology of host plants

The adaptation of three allopatric populations of a generalist moth, Operophtera brumata (L.), on two tree species, Prunus padus (L.) and Quercus robur (L.) which represent the extremes of the timing of budburst in spring, was studied in Finland and Sweden. The synchrony of the hatching and budbursting was monitored, and its importance to dispersal and growth of larvae was assessed by rearing cohorts of larvae, whose hatching dates were manipulated, on both hosts. In addition, the realised heritability of the hatching time was estimated.Experimental populations hatched in approximate synchrony with the budburst of their original host species. As a result of the manipulation of the hatching dates of larvae, the growth rates of larvae decreased and the dispersal rates increased on both hosts in relation to the ageing of foliage. The realised heritability of hatching times was rather high (0.63). There were fewer differences in the host use efficiency and behaviour of the experimental populations than in the hatching phenology. The synchrony of hatching with the budburst of the local dominant host plant is probably a result of stabilising selection.     

Non-feeding by adult gynoparae of Rhopalosiphum padi and its bearing on the limiting resource in the production of sexual females in host alternating aphids

Radioactive plants were used to determine whether gynoparae of the bird cherry-oat aphid Rhopalosiphum padi (L.) fed after settling on Prunus padus (L.). Despite being starved for 24 h they did not feed. The adaptive significance of the short adult life and low fecundity of gynoparae was discussed in the light of these results.

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La consommation des synopares adultes de Rhopalosiphum padi et son importance sur la limitation des resources dans la production de femelles sexuées chez les pucerons à hôtes alternatifs

Résumé Des plantes radioactives ont été utilisées pour déterminer si les gynopares de Rhopalosiphum padi se nourrissaient après installation sur Prunus padus. Bien qu'affamés pendant 24 h, il ne se sont pas nourris. La signification adaptative de la brève vie des adultes et de la faible fécondité des gynopares a été examinée à la lumière de ces résultats.

     

Probing behavior of bird cherry-oat aphid Rhopalosiphum padi (L.) on native bird cherry Prunus padus L. and alien invasive black cherry Prunus serotina Erhr. in Europe and the role of cyanogenic glycosides

The probing behavior of bird cherry-oat aphid Rhopalosiphum padi was studied on its natural winter host in Europe, the bird cherry Prunus padus, and on the invasive black cherry Prunus serotina, on which spring generations of R. padi do not survive. The EPG-recorded behavior of R. padi on bird cherry and black cherry showed differences in crucial aspects of probing and feeding. The period of the pre-phloem penetration was twice as long and rarely interrupted in aphids on bird cherry as opposed to aphids on black cherry. On black cherry, there was a considerable delay between finding and accepting the phloem. Aphids that had sampled phloem sap either refused to ingest it or the ingestion periods were very short. Amygdalin and prunasin (cyanogenic glycosides present in leaves of Prunus) seriously impeded ingestion activities when applied in pure sucrose diet. The role of amygdalin and prunasin in winter host plant selection and host alternation in R. padi is discussed.     

Lack of physiological improvement in performance of Callosamia promethea larvae on local host plant favorites

Summary As a species, the promethea silkmoth, Callosamia promethea (Saturniidae: Lepidoptera) exhibits a wide host range on 6–10 families of plants, although specific populations are known to have local foodplant favorites. We tested the hypothesis that larvae from a particular host plant lineage would show physiological adaptations to this host compared with larvae from other host plant lineages. We found no evidence that larval survival and growth was any better for larvae fed the natural plant of the parental population than for larvae from other host lineages. These natural host lineages include: black cherry (Prunus serotina Ehrh.), tuliptree (Liriodendron tulipifera L.), sassafras (Sassafras albidum (Nutt.) Nees) and spicebush (Lindera benzoin (L.) Blume). The only apparent manifestation of physiological specialization was the inability of tuliptree lineages of C. promethea to survive on paper birch (Betula papyrifera Marsh), although this may reflect the geographical pattern of adaptation to birch, rather than a negative correlation with adaptation to tuliptree. These results suggest that for C. promethea larvae, growth performance and survival is primarily influenced by plant nutritional quality, rather than physiological adaptations to the locally preferred host plant.     

Reproduction and probing behaviour of the bird cherry‐oat aphid Rhopalosiphum padi on detached leaves and leaves on intact seedlings of the winter host,bird cherry Prunus padus

Video recording and electrical penetration graph techniques are used simultaneously to investigate host acceptance (in terms of reproduction) and probing activities of autumn migrants (gynoparae) of the bird cherry‐oat aphid Rhopalosiphum padi (L.) on detached leaves and on leaves on intact plants of the winter host, bird cherry Prunus padus. There are no significant differences between the times taken to first parturition or the number of nymphs produced over a 6‐ or 18‐h period on intact plants or detached leaves. Stylet probing activities (i.e. total probe duration, xylem ingestion, the time to first phloem contact and phloem salivation) are also similar in the two situations.     

Does a Specialist Parasitoid Adapt to its Host on a New Host Plant?

The host plant expansion of a diamondback moth, Plutella xylostella (L.) (DBM) strain to snowpea (Pisum sativum L.) raised the question whether a specialist parasitoid Diadegma semiclausum (DS) could be conditioned to locate and parasitize its host on the new host plant. In a specialist parasitoid a behavioural change towards a plant outside the normal host plant range of its host due to developmental experience is not expected. The responsive behaviour, parasitism rates and fitness of three subsequent DS generations were investigated on the snowpea-strain of DBM. After three generations of DS on the pea 62.5% of females chose an DBM-infested pea plant over DBM infested cabbage. Only 16.4% of cabbage-reared DS was attracted to infested pea. Rearing of the parasitoid in host larvae on peas significantly increased the number of larvae parasitized on this host plant in the first generation; however, there was no further increase in generations 2 and 3. Larval mortality was similar for all parasitoid/DBM combinations on both host plants, but significantly higher mortality occurred in parasitoid pupae from peas. Development time of the parasitoid was slightly prolonged on the pea strain of DBM. The number of females produced by parasitoids reared on the pea strain of DBM was significantly reduced as compared to D. semiclausum reared on the cabbage strain on both host strains. Results show that DS has the potential to change its responsive behaviour in order to locate its host on a new host plant. According to the current view, a specialist parasitoid is not expected to change its reaction to a plant outside the normal host plant range of its host. Within 3 generations, responsive behaviour towards snowpea could be increased. However, fitness trade-offs, especially an extreme shift in sex ratio to males reduced reproductive success.     

Monilinia ssiori sp. nov. in the Sclerotiniaceae, causing leaf blight and young fruit rot of Prunus ssiori in Japan

The leaf blight and mummy fruit disease fungus of Prunus ssiori in northern Japan is newly named M. ssiori, as a fourth member in the M. padi group of section Disjunctoriae of the genus Monilinia (Sclerotiniaceae). It has been misidentified with Monilinia kusanoi, but recent studies show it is different from M. kusanoi as well as other related species on prunaceous hosts in respect to host relation, pathogenicity, morphology, and gene analysis.     

Host finding in the pine weevil Hylobius abietis: effects of conifer volatiles and added limonene

The attraction of Hylobius abietis (L.) (Coleoptera: Curculionidae) to -pinene was recently shown to be inhibited by limonene. This study shows that added limonene also affects the response of H. abietis to odours from natural host material. The experiments included both pieces of Scots pine suspended in pitfall traps and planted pine seedlings that had either been manually wounded or left intact. By adding a limonene dispenser, the catch of pine weevils in pine-baited traps was reduced by about 80–90% in spring and autumn and by about 40–50% in summer. Limonene substantially reduced the rate at which manually wounded seedlings were attacked by weevils in spring and autumn. For intact seedlings, a limonene-induced reduction in attack rate was found only in autumn. In the absence of limonene, a small wound made on the stem of a seedling increased its probability of being attacked by about fivefold. This figure was estimated by comparing data from all experimental periods with a model assuming a constant ratio of attack rates for the two kinds of seedlings.     

Influence of endophyte infection, plant age and harvest interval on Rhopalosiphum padi survival and its relation to quantity of N-formyl and N-acetyl loline in tall fescue

Rhopalosiphum padi L. (Homoptera: Aphididae) is sensitive to loline alkaloids present in tall fescue, Festuca arundinacea Shreb., infected with the endophytic fungus, Acremonium coenophialum Morgan-Jones & Gams. Aphid survival was higher on endophyte-free plants regardless of plant age after germination or age of regrowth tissue after clipping. Survival of aphids on endophyte-infected grass was lower on young tissue but increased as plants aged, although it never reached the same level on endophyte-free plants. Both N-formyl and N-acetyl loline increased as uncut or regrowth tissue aged; however, this was influenced by the age of the plant at the initial cut and the clipping frequency. Although even small amounts of loline cause high aphid mortality, the aphids are able to survive on endophyte-infected plants if the tillers have senescing leaves which contain lower amounts of loline. Preference for senescing leaves may help R. padi avoid plant parts containing high amounts of toxic allelochemicals, thus contributing to higher numbers of aphids on older, endophyte-infected plants.     

Are endophyte-mediated effects on herbivores conditional on soil nutrients?

Neotyphodium endophytes are assumed to have mutualistic relationship with their grass hosts, mainly resulting from mycotoxin production increasing plant resistance to herbivores by the fungus that subsists on the plant. To study importance of often ignored environmental effects on these associations, we performed a greenhouse experiment to examine the significance of endophyte infection and nutrient availability for bird-cherry aphid (Rhopalosiphum padi) performance on meadow fescue (Lolium pratense). Naturally endophyte-infected (E+), uninfected (E–), or manipulatively endophyte-free (ME–) half-sib families of meadow fescue were grown on two soil nutrient levels. Endophyte infection reduced aphid performance in general. However, to our knowledge, this is the first study to demonstrate experimentally that herbivore performance decreases on E+ host plants with increasing availability of nutrients in soils. Potential improvement in herbivore performance in high nutrient soils and decreased plant performance in low nutrient soils in ME– plants, compared to E– and E+ plants, suggests that loss of endophyte infection after long coevolutionary relationship may be critical to plant fitness.     

Host and site selection by larval stages and adults of the parasitic copepod Lernaeenicus sprattae (Sowerby) (Copepoda,Pennellidae) in the Oslofjord

Sprat caught by beach seining during the period 1977–1980 were examined for adult Lernaeenicus, and both sprat and herring caught in 1981–1983 were examined for all parasitic stages. Lernaeenicus was never found on herring. Adult L. sprattae were always found in the sprat's eye, usually (89%) in the upper rear quadrant of the eye. Double and triple infection of one or both eyes were found. Infection experiments support results from field work. The sprat is the only intermediate and final host of L. sprattae. In laboratory experiments, copepodites showed no preference for particular size groups of sprat. The infective stages, copepodid and adolescent female, are both initially randomly distributed on the host before the copepodites move towards the fins and the gravid females to the eye.     

Breeding biology of Tenagomysis tasmaniae Fenton,Anisomysis mixta australis (Zimmer) and Paramesopodopsis rufa Fenton from south-eastern Tasmania (Crustacea: Mysidacea)

The number, size and developmental stage of young in the brood pouch of female Tenagomysis tasmaniae, Anisomysis mixta australis and Paramesopodopsis rufa was recorded throughout the year. Breeding was intensive from spring till the end of autumn for the three species. Calculation of the egg ratio for each species showed that their major reproductive peaks occurred during spring and summer. A winter depression in the breeding cycle was observed for T. tasmaniae and P. rufa, but A. mixta australis ceased breeding during winter. Seasonal variation in the length of gravid females and number of young carried was evident for these three species. Females were longer in spring and summer and carried more young than in autumn and winter. A linear relationship between female length and brood size was demonstrated for each species; annual and seasonal equations were calculated for females carrying each developmental stage. The seasonal equations showed that for a female of given length fecundity was greater during spring than any other season. Natality was estimated to be highest during late spring, summer and early autumn for the three species. No seasonal variation in the size of eggs was evident for the three species. The reproduction pattern of T. tasmaniae, A. mixta australis and P. rufa appears to be very similar to that reported for the majority of iteroparous coastal temperate mysids throughout the world.