Sub-lethal effects of deltamethrin residues on the within-crop behaviour and distribution ofCoccinella septempunctata

The behaviour and distribution of adultCoccinella septempunctata L. (Coleoptera: Coccinellidae) were recorded in two plots of winter wheat infested with the cereal aphidsSitobion avenae (F.) andMetopolophium dirhodum (Walk.) (Homoptera: Aphididae). One plot was sprayed with the pyrethroid insecticide deltamethrin at a rate of 6.25 g a.i./ha and the other was left unsprayed. Single ladybird beetles were released sequentially on the ground at the centre of the sprayed and unsprayed plots and their behaviour and position in the crop canopy were recorded at 30 second intervals for a total of 15 min per beetle. Assessments, with fresh beetles, continued for four days after spray application with a total of eighty ladybird beetles observed. The 15 min period was selected to avoid lethal effects and no ladybird beetles were killed or knocked down as a result of exposure to deltamethrin residues during this period. Significant differences were found between the overall behaviour patterns ofC. septempunctata in the untreated and deltamethrin treated plots up to three days after the spray application. Ladybird beetles exposed to deltamethrin residues were observed to walk and groom significantly more frequently and to rest significantly less frequently than those in the unsprayed plot. Significant differences were also found between the observed distribution of ladybird beetles in the sprayed and unsprayed crop canopies, with higher numbers of observations towards the bottom of the crop canopy and on the ground in the deltamethrin treated plot than in the untreated plot during the first two days after deltamethrin application. Upon the foliage itself, ladybird beetles were observed significantly more frequently on the abaxial leaf surface in the deltamethrin treated crop compared with the untreated crop. The results are discussed in terms of possible evidence for the repellency of deltamethrin toC. septempunctata and also the implications for integrated pest management of changes in predator behaviour and crop distribution resulting from sub-lethal uptake of insecticides.     

Effects of honeydew and insecticide residues on the distribution of foraging aphid parasitoids under glasshouse and field conditions

The opposing effects of attraction to host-derived kairomones and repellency from the pyrethroid insecticide deltamethrin were investigated with aphid parasitoids from the genus Aphidius (Hymenoptera: Aphidiinae). The spatial distribution of female parasitoids was recorded in a series of experiments conducted in a small glasshouse containing wheat plants either infested with cereal aphids, Sitobion avenae (F.) (Homoptera: Aphididae), uninfested or treated with the recommended field concentration of deltamethrin. The number of parasitoids per plant were counted at 0.5 h, 1 h and then at one hourly intervals up to 8 h after release. Parasitoids showed a strong aggregation response to aphid-infested plants compared to adjacent uninfested plants. With the introduction of insecticidetreated plants around the aphid-infested plants, parasitoids showed a greater tendency to disperse away, resulting in fewer parasitoids on plants and significantly lower rates of aphid parasitism. The degree of aphid fall-off from plants was a good indicator of parasitoid foraging activity. In field studies, using sticky traps to measure the activity of parasitoids in plots sprayed with water, deltamethrin and/or an artificial honeydew solution, repellent properties were evident for up to 2 days after application. The attraction/arrestment stimuli associated with the honeydew solution were sufficient for parasitoids to continue searching insecticide-treated areas. The implications of these findings for parasitoids searching crops contaminated with aphid-derived kairomones and insecticides are discussed.     

Mobility of apterous grain aphids Sitobion avenae within wheat fields

In a field study of the grain aphid Sitobion avenae (F.) almost all apterous aphids were found to move from the natal ear before reproducing, and alatiform nymphs usually moved between ears during their development. Apterae were very mobile, with over 50% staying on one ear for less than 24 h. Movements did not appear to be a response to climatic conditions, predation or colony density.

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Résumé Lors de l'étude de Sitobion avenae dans la nature, on a constaté que presque tous les individus aptères avaient quitté leur épi d'origine avant la reproduction, et que les larves de type ailé s'étaient déplacées généralement entre les épis pendant leur développement. Les aptères étaient très mobiles, la majorité restant moins de 24 h sur le même épi. Ces déplacements n'ont pas semblé dépendre des conditions climatiques, de la prédation ou de la densité des colonies.

     

Toxicity of the new pyrethroid insecticide,deltamethrin, to Daphnia magna

The toxicity of deltamethrin, a synthetic pyrethroid insecticide, was determined under standardized conditions (ISO, 1982) in neonates and juveniles of Daphnia magna. Neonates (6 to 24 h old) were more sensitive than juveniles (48 to 72 h old). The 24- and 48-h EC₅₀s (immobilization) in neonates were 0.113 and 0.031 µg l^–1, respectively. The toxicity of deltamethrin was highly toxic. The 96-h EC₅₀ was in the ppt (µg l^–1) range. Toxicity tests with Daphnia may be used to detect toxic residues in water and sediment in areas treated with deltamethrin and other highly toxic pyrethroid pesticides.     

The spatial and temporal distribution of the grain aphid Sitobion avenae in winter wheat

The spatial and temporal distribution of the grain aphid Sitobion avenae F. (Homoptera: Aphididae) was studied within a field of winter wheat during the summer of 1996. Sampling was done using four nested grids comprising 133 locations. Analysis by Taylor's power law gave results typical for insect populations. Analysis by SADIE (Spatial Analysis by Distance Indices) showed spatial pattern due to edge effects and sampling scale, and positive but mild spatial association, although spatial patterns were ephemeral. Reasons for these findings and the implications for integrated crop management are discussed.     

The influence of insecticide residues on primary parasitoid and hyperparasitoid foraging behaviour in the laboratory

The behaviour of Aphidius rhopalosiphi DeStefani-Perez (Hymenoptera: Braconidae) was recorded in laboratory bioassays in the presence and absence of both residues of honeydew from the English grain aphid, Sitobion avenae (F.) and the pyrethroid insecticide, deltamethrin. Insecticide concentrations ranged from the field recommended dose rate (6.25 g ai/ha in 2001 water) to 1/16th of field rate. Parasitoids responded strongly to patches of honeydew on filter papers, however the addition of increasing concentrations of deltamethrin caused increasingly early departure from the honeydew-treated areas. Parasitoids pre-exposed to field concentration residues for between 1 and 20 min showed shorter retention times and abnormal types of behaviour on honeydew-treated patches compared to control wasps. Recovery of normal behaviour patterns occurred over a 12 h period away from the insecticide source. Further laboratory studies examined the foraging of A. rhopalosiphi and one of its associated hyperparasitoids, Dendrocerus carpenteri (Curtis), on mature wheat plants treated with an artificial honeydew solution and deltamethrin. Wasps exposed to deltamethrin residues were observed to spend shorter visit times, to groom more frequently and to rest less frequently than those on insecticide-free plants. Differences were found between the distribution of parasitoids on insecticide-treated and untreated plants, with D. carpenteri showing greater movement down insecticide-treated plants accompanied with an increase in time spent on abaxial leaf surfaces compared to A. rhopalosiphi. The results are discussed in terms of the repellency of deltamethrin and the implications for integrated pest management of differing sub-lethal insecticide effects on primary parasitoid and hyperparasitoid behaviour.     

The potential of predators in natural control of aphids in wheat: Results of a ten-year field study in two German landscapes

In the present study, we investigated the natural control of aphids by predators in wheat fields in a low (L) and high-input cropping region (H) of Germany during a 10-year period. Data for the statistical analyses were obtained from weekly after the start of aphid emergence. The mean annual aphid indices, calculated as the sum of Sitobion avenae (Fabr.), Rhopalosiphum padi (L.), Metopolophium dirhodum (Walk.)(Homoptera: Aphididae), were 30.4 and 81.5 × 103 aphid days per m2, for L and H, respectively. Nine predator fractions were analysed: Coccinella septempunctata L., adults (1) and larvae (2), Propylea quatuordecimpunctata (L.) (Coleoptera: Coccinellidae) adults (3) and larvae (4), syrphid larvae (mostly Episyrphus balteatus [De Geer] (Diptera: Syrphidae)) (5), Chrysoperla carnea Steph. (Neuroptera: Chrysopidae) larvae (6), and adult carabids (7), staphylinids (8) and spiders (9). The two sites were comparable in terms of the mean size of the overall predator community, expressed in predator units (PU): 4.9 PU/m2 (L) vs. 5.4 PU/m2 (H). Most predator fractions responded numerically to increasing aphid densities. The numerical response was strongest in syrphid larvae, scarcely detectable in adult coccinellids, and virtually non-existent in epigeic arthropods. Multiple regression models revealed indirect relationships between the weekly overall predator community densities (PU/m2) and individual predator fractions (individuals/m2) and absolute rates of aphid density increase (individuals/m2) one or two weeks after baseline. A site-independent reduction of the aphid density increase to nil (y = 0) was observed at 3.9 to 4.2 PU/m2. Consequently, the 2.7 times higher aphid density at H cannot be attributed to the presence of fewer predators or lower effects of the overall predator community or of any individual predator fraction.     

Populations ofSitobion avenae andAphidius ervi on spring wheat in the northwestern United States Spatial distribution and sequential sampling plans based on numerical and binomial counts

During a three-year field survey of the English grain aphid,Sitobion avenae (F.) (Homoptera: Aphididae) and the parasitoid,Aphidius ervi Haliday (Hymenoptera: Aphidiidae; represented by aphid mummies) on spring wheat, data sets were generated which consisted of 47 estimates of mean density (m aphids per tiller), variance (s ²), and the proportion of empty tillers (P ₀ forS. avenae and 22 estimates of the same statistics forA. ervi, respectively. Each estimate of the aphid and parasitoid populations was based on counts per individual tiller on each sampling occasion. Taylor's power law was found to fit the data sets better than Iwao'sm ^*−m regression model. Taylor's slope forS. avenae (1.3076) was significantly higher than that forA. ervi (1.1519), indicating thatS. avenae was more aggregated thanA. ervi. Based on temporal changes in spatial aggregation represented by an index, 1/k [wherek was estimated asm ²/(s²−m)],S. avenae was most aggregated at low densities during the early infestation period and tended to be less aggregated as density increased, whereasA. ervi was more likely to be randomly distributed. A commonk was detected for neitherS. avenae norA. ervi because the slopes from the linear regression ofk onm significantly exceeded 0. Sequential sampling plans, based on numerical (complete) and binomial (presence or absence) counts, for the aphid and parasitoid species were developed using Taylor's parameters and those estimated from the linear regression of ln(m) on ln[−ln[P ₀], respectively. Suggestions are made concerning the use of the sampling plans and the levels of precision that may be attained.     

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.

     

Interspecific interactions in a tri-trophic arthropod system: effects of a spider on the survival of larvae of three predatory ladybirds in relation to aphids

The effects of the crab spider, Misumenops tricuspidatus (Fabricius), on the larval survival of three ladybird species, Harmonia axyridis Pallas, Coccinella septempunctata L., and Propylea japonica L., in relation to aphids were investigated in the laboratory. Predation by the spider on the three ladybird species differed. All the larvae of C. septempunctata, none of H. axyridis, and an intermediate number of P. japonica were attacked and eaten by the spider. All the larvae of H. axyridis suffered mortality due to cannibalism or starvation in the treatments with and without a spider. In case of C. septempunctata, however, mortality in the early instars was significantly greater in the treatment with a spider than without a spider and no larvae developed into pupae due to predation. In the treatment without a spider, the majority of the larvae in the former treatment suffered mortality due to cannibalism or starvation, and only 13.3% of larvae developed into the adult stage. In the case of P. japonica, mortality was mainly attributed to predation in the treatment with a spider and only 26.7% became adult. In comparison, 86.7% of larvae survived to the adult stage in the treatment without a spider. In addition, in both H. axyridis and C. septempunctata, the development of young larvae was significantly slower in the presence of a spider, but this was not the case with the older larvae of H. axyridis, which indicates that the effect of the spider on larval development changed with the developmental stage of the larvae in this species. However, the spider had no significant effect on the developmental time of P. japonica larvae. Although both the spider and the ladybirds significantly affected the number of aphids, they did not have an additive effect on aphid abundance. The interactions between the spider and the ladybirds, such as intraguild predation or competition, caused them to reduce aphid population density less than the ladybirds did on their own. The effect of the spider on the larval performance of three predatory ladybirds was found to be unequal in terms of their vulnerability to predation and rate of larval development and it depended on the species and developmental stage of the ladybird.     

Production of sexual morphs by the monoecious cereal aphid Sitobion avenae

The effects of photoperiod and temperature on the production of sexual forms by two clones of Sitobion avenae, the grain aphid, were examined. One clone did not produce sexual forms, whereas the other did under conditions of short light (<14 h) and low temperature (15°C). Temperature and photoperiod interacted to some extent both in the production of oviparae and of males. Even when the sexual forms were produced there was always a proportion of virginoparae.

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Production de sexués par Sitobion avenae, puceron monoécique

Résumé Un clone de Sitobion avenae a produit sans difficulté jusqu'à 7 générations de sexués en réaction à des photopériodes courtes (<14 h) et à de faibles températures (15°C) tandis qu'un autre clone n'en produit aucune.La proportion de sexués différait suivant les parents. La production de mâles avait une nette tendance à apparaître lors des reproduction les plus tardives. Dans les générations tardives avec jours courts, les vivipares (virginopares et gynopares) avaient eu tendance à être produits à la fin de la période de reproduction.Les gynopares (c'est à dire les parents d'ovipares) de S. avenae étaient surtout aptères, mais comprenaient beaucoup plus d'ailés que les virginopares obtenus dans les mêmes conditions. Un vivipare était induit comme gynopare (ou ses embryons étaient déterminés comme ovipares) avant la naissance, mais cette détermination pouvait apparemment encore être inversée en soumettant l'insecte à de longues photopériodes et à de hautes températures jusqu'à deux jours après la naissance.Aucun S. avenae ovipare n'a été produit jusqu'à la troisième génération. Aucun ovipare n'a été produit avec des photopériodes supérieures à 13 H 30 à 10°C, 13 H à 15°C et 8 H à 20°C. La proportion d'individus produisant des ovipares à 15°C a été plus faible qu'à 10°C pour toutes les photopériodes, et à cette dernière température beaucoup plus de vivipares étaient gynopares que virginopares.Les basses températures ont été vraisemblablement le facteur dominant de stimulation de la production de mâles de S. avenae, mais cependant il semble qu'un plus grand nombre de mâles a été produit aux températures et photopériodes intermédiaires qu'aux extrêmes.La capture de mâles ailés de S. avenae dans des pièges à succion a été généralement limitée à Octobre-soit à peu prés le moment prévu par les expériences de laboratoire. Des mâles de S. avenae sont aussi capturés fréquemment et été, ce qui peut être lié à des hivers précédents doux.

     

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.

     

Interaction between zinc nutritional status of cereals and Rhizoctonia root rot severity

An inverse correlation between plant Zn concentration and the severity of Rhizoctonia root rot, described in an earlier paper, was examined in two experiments in a growth chamber. In the first experiment, wheat (Triticum aestivum cv Songlen) was planted in a Zn deficient soil with and without added Zn, and combined factorially with different inoculum densities of Rhizoctonia solani anastomosis group 8. When Zn was added, the percentage of seminal roots infected with R. solani was significantly lower compared to the treatments without added Zn, showing that low Zn potentiated the disease. A subsequent factorial experiment of four inoculum densities and six Zn levels, (0, 0.01, 0.04, 0.1, 0.4 and 6.0 mg Zn kg^–1 soil) was conducted to investigate the Zn effect in more detail. Disease severity was markedly decreased by the higher Zn applications; the disease score dropped sharply between treatments of Zn_0.04 and Zn_0.1, a difference which was reflected in the plant yield response to Zn. For both experiments the Zn concentrations in shoots were significantly different only among Zn treatments, not among the inoculum treatments. This indicated that inoculum density or disease severity did not reduce Zn concentration in the plant. Thus, disease did not exaggerate Zn deficiency, but rather, Zn sufficiency suppressed disease severity. A potentiating link between Zn nutrition and disease severity is thereby established, although this type of experiment did not indicate the mechanism of the Zn effect.     

Negative dietary effects of Colorado potato beetle eggs for the larvae of native and introduced ladybird beetles

Aphidophagous ladybird beetles (Coleoptera: Coccinellidae) are attracted to and feed heavily on aphids, but many species will also feed opportunistically on other prey that they encounter. In potatoes (Solanum tuberosum L.) in Washington State, USA, coccinellids feed on both green peach aphids (“GPA,” Myzus persicae Sulzer) and eggs of the Colorado potato beetle (“CPB,” Leptinotarsa decemlineata Say). The guild of aphidophagous ladybirds includes two native species, Hippodamia convergens Guérin-Méneville and Coccinella transversoguttata Brown. Recently, an introduced species, Coccinella septempunctata L., has invaded and apparently displaced its native congener. A second exotic, Harmonia axyridis Pallas, has colonized the area and is becoming more abundant. We compared larval development of each species on a monotypic diet of GPA, a monotypic diet of CPB eggs, or a mixed diet of both GPA and CPB eggs. Our goal was to answer two questions: (1) do larvae of the four ladybird species benefit from including CPB eggs in their diet and (2) do the four ladybird species differ in their ability to utilize CPB eggs as prey? No larva of any species completed development on a pure diet of CPB eggs, and survivorship was highest for all species when they fed on a pure diet of GPA. One native species, H. convergens, and one exotic species, H. axyridis, exhibited significantly lower survivorship on a mixed diet of both CPB eggs and GPA, compared to a pure GPA diet; H. axyridis also took longer to develop from egg to adult when both prey were provided. Survivorship of the two Coccinella spp. was not altered by the inclusion of CPB eggs with GPA, although CPB eggs lengthened the development time of C. transversoguttata. Adult size was not consistently affected by diet for any of the coccinellids. Overall, no ladybird species benefited from the inclusion of potato beetle eggs in its diet. The two Coccinella species responded similarly to the inclusion of CPB eggs, and so we would not expect any difference in the success of coccinellid larval development in potato fields following the replacement of C. transversoguttata by C. septempunctata. Hippodamia convergens and H. axyridis, the two species whose survivorship was depressed by combining CPB egg and aphid prey, were also the two species that consumed the greatest number of CPB eggs during successful larval development. A comparison of total egg consumption by each species cohort suggested that displacement of the other species by H. axyridis would not alter CPB biological control, because the higher per capita feeding rate by H. axyridis larvae compensated for individuals’ greater mortality risk on a diet including CPB eggs.     

The cost of switching from asexual to sexual reproduction in an aphid

The fecundity of asexual and sexual morphs of Sitobion avenae (F.) is compared. The cost of sex is quantified in terms of the reduction in numbers of offspring produced. The cost is much greater than the halving of fecundity that is usually assumed to be associated with the production of males.

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Résumé Les fécondités des virginipares ailés, des gynopares ailés et des ovipares d'un même clone de Sitobion avenae ont été déterminées à 12°C. Bien que les types qui ayant donné naissance à des sexués aient eu une fécondité initiale plus élevée que celle de la lignée asexuée, celle-ci a été significativement plus féconde après 15 jours de reproduction.Les résultats du laboratoire ont été utilisés pour évaluer la fécondité de lignées sexuées et asexuées pendant quatre générations à 12°C. La reproduction parthénogénétique a présenté un avantage numérique sur la reproductions sexuée dans le rapport 320/1. La reproduction sexuée a été accompagnée d'une diminution significative de la fécondité potentielle. Bien que la reproduction sexuée de S. avenae soit avantageuse dans les régions où les hivers sont durs, parce que les oeufs sont résistants au froid, les lignées asexuées présentent un avantage considérable en ce qui concerne la fécondité potentielle pour l'augmentation de la population en automne et elles conservent de plus la capacité d'un accroissement numérique ultérieur si l'hiver est doux.

     

Methods of hatching the eggs and rearing the fundatrices of the English grain aphid, Sitobion avenae

Several methods for hatching the eggs and rearing individuals of the first generation (fundatrices) of Sitobion avenae were investigated. The most successful methods were incubation of the eggs on grass seedlings at 2°C and rearing the fundatrices on grass seedlings (overall survival 66%) and incubation of the eggs in plastic boxes at 2°C and rearing the fundatrices on wheat seedlings (overall survival 62%).

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Résumé L'éclosion des oeufs de S. avenae peut être induite par le transfert à 10°C ou 12°C, après une incubation de 75–120 jours à 2°C. Le pourcentage le plus élevé d'éclosions a été obtenu quand les oeufs avaient incubé pendant 100 à 110 jours à 2°C (67% at 71.5% respectivement) dans des petites boîtes de plastique, ou pendant 100 jours à 2°C sur des pousses de graminées (73.5%). Si les oeufs sont pondus sur blé, la plante ne peut pas tolérer la période d'incubation, mais cet obstacle peut être surmonté en obligeant les ovipares à pondre leurs oeufs sur de pousses de graminées, comme Poa annua, hôte convenable pour les fondatrices. Les ovipares peuvent aussi pondre sans difficultés sur autre chose que des végétaux, et des récipients peuvent ètre mis à incuber sans contenir du matériel végétal.

     

Low field performance of an aphid parasitoid,Aphelinus abdominalis,efficient in the laboratory [Hym., Aphelinidae]

Life history traits of two inbred lines of the aphid parasitoidAphelinus abdominalis (Dalman) were evaluated in the laboratory. Both lines exhibited high fecundities, produced a high proportion of female offspring and showed to be long-lived. However, release experiments in winter wheat fields failed to achieve establishment of the parasitoids. In addition, the wasps parasitized considerably fewer aphids in two types of field cages than they did in the laboratory. AsA. abdominalis eggs and/or larvae arrested in development were not found in dissected aphids from field cages, we conclude that the wasps were unable or unwilling to parasitize high numbers of hosts in the cages. The reason for this odd behaviour is unknown.     

Adaptations of Coccinella magnifica Redtenbacher, a Myrmecophilous Coccinellid, to Aggression by Wood Ants (Formica rufa Group). I. Adult Behavioral Adaptation, Its Ecological Context and Evolution

Coccinella magnifica is an obligate associate of Formica rufa-group ants. The congener Coccinella septempunctata can serve as a model for its nonmyrmecophilous ancestor. Formica rufa behavior toward these two ladybirds, and their behavior, was compared. Although C. magnifica was rarely attacked on ant trails, it was usually attacked on tended aphid colonies. Coccinella septempunctata was more readily attacked. The two ladybirds' behavior was similar on trails, but C. magnifica used more defense and C. septempunctata more rapid escape behavior on aphid colonies. Only C. magnifica fed upon tended aphids. Chemical adaptation to overcome ant aggression probably exists in C. magnifica, but it possesses almost no novel behaviors to counter ant aggression. Instead, modifications have occurred in the expression of behaviors present in C. septempunctata.     

Gas exchange of ears of cereals in response to carbon dioxide and light

One cultivar each of spring wheat (Triticum aestivum L. cv. Arkas), oat (Avena sativa L. cv. Lorenz), and barley (Hordeum vulgare L. cv. Aramir) was chosen in order to study the relative contributions of individual bracts to the gas exchange of whole ears. The distribution and frequency of the stomata on the bracts were examined. Gas exchange was measured at normal atmospheric CO₂ (330 bar) and at high CO₂ (2000 bar) on intact ears and on ears from which glumes or lemmas and pleae (wheat and oat) or awns (barley) had been removed.The relative contribution to the gas exchange of the whole organ is highest for the awns of barley ears. In wheat, the contribution of the glumes is slightly higher than that of the inner bracts before anthesis. Two weeks after anthesis the inner bracts contribute more than the glumes. This tendency of increasing importance of the inner bracts is also found in oat ears, but the relative amount of CO₂ uptake by the glumes is higher than in wheat. These changes during ontogeny result from the better supply of light to the inner bracts caused by opening of the ears' structures during grain filling, which in part compensates for the decreasing photosynthetic capacity.The ratio of the photosynthesis rate at high CO₂ to that at normal CO₂ is lower for the glumes of oat and for the awns of barley than for the other bracts.Abbreviations A³³⁰, A²⁰⁰⁰ net photosynthesis rate, A³³⁰ at normal atmospheric CO₂ (330 bar), A²⁰⁰⁰ at high CO₂ (2000 bar) - PPFD photosynthetic photon flux density - p^c intercellular partial pressure of CO₂