Intraguild Interactions between the Mealybug Predators Cryptolaemus montrouzieri and Chrysoperla carnea.

The ladybird Cryptolaemus montrouzieri and the green lacewing Chrysoperla carnea have shown potential for use in augmentative biological control of mealybug pests in greenhouse crops. In the context of combining these predators within an integrated pest management system, the risk of negative intraguild interactions between both predators was evaluated in a laboratory setting. Different life stages of either predator were confronted in petri dish arenas containing a Ficus benjamina leaf, and after 24 h the incidence and direction of intraguild predation (IGP) was recorded for each combination. The effect of adding Planococcus citri nymphs or Ephestia kuehniella eggs as extraguild prey on the level of IGP was also studied. IGP was frequently observed between the two predator species and was asymmetrical in favour of C. carnea in most cases. The presence of extraguild prey reduced the number of IGP events between the predators to a similar extent. The relevance of the observed intraguild interactions for the combined use of these predators in protected cultivation is discussed.

Cold-born killers: exploiting temperature-size rule enhances predation capacity of a predatory mite.

BACKGROUND: The temperature-size rule is a well-known example of phenotypic plasticity in ectothermic organisms. When exposed to colder temperatures, ectotherms develop more slowly, but mature at larger body sizes and vice versa at higher temperatures. We investigated whether a phytoseiid predatory mite can obtain a larger body size by rearing it at a low temperature and how the increased body size affected predatory performance on its natural prey. Therefore, we allowed the predatory mite Amblydromalus limonicus (Garman & McGregor) (Acari: Phytoseiidae) to develop at either 15 or 25 °C. RESULTS: A. limonicus reared at 15 °C had a 6% larger body size than those reared at 25 °C. Larger predators showed higher predation rates on first instars of the western flower thrips, Frankliniella occidentalis Pergande (Thysanoptera: Thripidae), with 9.6 instars/female/day and 8.5 instars/female/day, for larger and standard-sized females, respectively. After three generations reared at 15 °C, body size did not increase any further. When reared for five generations at 15 °C, larger A. limonicus females demonstrated a better ability to subdue second-instar F. occidentalis. CONCLUSION: Low juvenile rearing temperatures may result in phytoseiid predators with a predator/prey size benefit that could improve their biological control function. © 2019 Society of Chemical Industry.

Adventitious rooting of Chrysanthemum is stimulated by a low red:far-red ratio.

Adventitious rooting, a critical process in the vegetative propagation of many ornamentals, can be affected by both light intensity and light quality. We investigated the use of spectral light quality to improve adventitious rooting of Chrysanthemum morifolium cuttings by applying different combinations of blue, red and far-red light. Additionally, unrooted cuttings were treated before planting with two auxin transport inhibitors (TIBA and NPA) to study the effect of light quality on auxin biosynthesis and/or transport. Results showed that lowering the R:FR ratio (decreasing the phytochrome photostationary state, PSS) improved rooting significantly and decreased the inhibiting effect of the auxin transport inhibitor NPA. An extra decrease of PSS by adding blue light to a red + far-red spectrum further enhanced rooting. In contrast, adding blue light to solely red light decreased rooting, an effect which was more pronounced in combination with the auxin transport inhibitors TIBA and NPA. Our results show that phytochrome plays a role in adventitious root formation through the action of auxin, but that also blue light receptors interact in this process.

Heritability of symbiont density reveals distinct regulatory mechanisms in a tripartite symbiosis.

Beneficial eukaryotic-bacterial partnerships are integral to animal and plant evolution. Understanding the density regulation mechanisms behind bacterial symbiosis is essential to elucidating the functional balance between hosts and symbionts. Citrus mealybugs, Planococcus citri (Risso), present an excellent model system for investigating the mechanisms of symbiont density regulation. They contain two obligate nutritional symbionts, Moranella endobia, which resides inside Tremblaya princeps, which has been maternally transmitted for 100-200 million years. We investigate whether host genotype may influence symbiont density by crossing mealybugs from two inbred laboratory-reared populations that differ substantially in their symbiont density to create hybrids. The density of the M. endobia symbiont in the hybrid hosts matched that of the maternal parent population, in keeping with density being determined either by the symbiont or the maternal genotype. However, the density of the T. princeps symbiont was influenced by the paternal host genotype. The greater dependency of T. princeps on its host may be due to its highly reduced genome. The decoupling of T. princeps and M. endobia densities, in spite of their intimate association, suggests that distinct regulatory mechanisms can be at work in symbiotic partnerships, even when they are obligate and mutualistic.

Suboptimal Light Conditions Influence Source-Sink Metabolism during Flowering.

Reliance on carbohydrates during flower forcing was investigated in one early and one late flowering cultivar of azalea (Rhododendron simsii hybrids). Carbohydrate accumulation, invertase activity, and expression of a purported sucrose synthase gene (RsSUS) was monitored during flower forcing under suboptimal (natural) and optimal (supplemental light) light conditions, after a cold treatment (7°C + dark) to break flower bud dormancy. Post-production sucrose metabolism and flowering quality was also assessed. Glucose and fructose concentrations and invertase activity increased in petals during flowering, while sucrose decreased. In suboptimal light conditions RsSUS expression in leaves increased as compared to optimal light conditions, indicating that plants in suboptimal light conditions have a strong demand for carbohydrates. However, carbohydrates in leaves were markedly lower in suboptimal light conditions compared to optimal light conditions. This resulted in poor flowering of plants in suboptimal light conditions. Post-production flowering relied on the stored leaf carbon, which could be accumulated under optimal light conditions in the greenhouse. These results show that flower opening in azalea relies on carbohydrates imported from leaves and is source-limiting under suboptimal light conditions.

Supplemental food for Amblyseius swirskii in the control of thrips: feeding friend or foe?

BACKGROUND: In integrated pest management systems in greenhouse crops, the predatory mite Amblyseius swirskii is becoming increasingly important as a biological control agent of various pests, especially thrips and whiteflies. An emerging strategy to promote the predator's establishment and retention in the crop consists in providing food supplements. However, when faced with omnivorous pests, such as the western flower thrips, Frankliniella occidentalis, food supplements need to be applied with extreme care, in order not to boost population growth of the pest. This laboratory study was conducted to evaluate the impact of commercial products of Typha angustifolia pollen and decapsulated brine shrimp cysts (Artemia sp.) on populations of both pest and predator and on predator-prey interactions. RESULTS: Pollen was highly supportive for both F. occidentalis and A. swirskii, whereas Artemia cysts supported thrips populations to a lesser extent than those of the predator. Furthermore, a less pronounced reduction in thrips consumption by A. swirskii was observed in the presence of Artemia cysts as compared with T. angustifolia pollen. CONCLUSION: Artemia might be a valuable alternative to pollen for supporting populations of A. swirskii in order to improve thrips management, as they are less beneficial for the pest but do support population growth of A. swirskii.

Diurnal temperature variations affect development of a herbivorous arthropod pest and its predators.

The impact of daily temperature variations on arthropod life history remains woefully understudied compared to the large body of research that has been carried out on the effects of constant temperatures. However, diurnal varying temperature regimes more commonly represent the environment in which most organisms thrive. Such varying temperature regimes have been demonstrated to substantially affect development and reproduction of ectothermic organisms, generally in accordance with Jensen's inequality. In the present study we evaluated the impact of temperature alternations at 4 amplitudes (DTR0, +5, +10 and +15°C) on the developmental rate of the predatory mites Phytoseiulus persimilis Athias-Henriot and Neoseiulus californicus McGregor (Acari: Phytoseiidae) and their natural prey, the two-spotted spider mite Tetranychus urticae Koch (Acari: Tetranychidae). We have modelled their developmental rates as a function of temperature using both linear and nonlinear models. Diurnally alternating temperatures resulted in a faster development in the lower temperature range as compared to their corresponding mean constant temperatures, whereas the opposite was observed in the higher temperature range. Our results indicate that Jensen's inequality does not suffice to fully explain the differences in developmental rates at constant and alternating temperatures, suggesting additional physiological responses play a role. It is concluded that diurnal temperature range should not be ignored and should be incorporated in predictive models on the phenology of arthropod pests and their natural enemies and their performance in biological control programmes.

Alternating temperatures affect life table parameters of Phytoseiulus persimilis, Neoseiulus californicus (Acari: Phytoseiidae) and their prey Tetranychus urticae (Acari: Tetranychidae).

Increasing energy costs force glasshouse growers to switch to energy saving strategies. In the temperature integration approach, considerable daily temperature variations are allowed, which not only have an important influence on plant growth but also on the development rate of arthropods in the crop. Therefore, we examined the influence of two constant temperature regimes (15 °C/15 °C and 20 °C/20 °C) and one alternating temperature regime (20 °C/5 °C, with an average of 15 °C) on life table parameters of Phytoseiulus persimilis and Neoseiulus californicus and their target pest, the two-spotted spider mite Tetranychus urticae at a 16:8 (L:D) h photoperiod and 65 ± 5 % RH. For females of both predatory mites the alternating temperature regime resulted in a 25-30 % shorter developmental time as compared to the corresponding mean constant temperature regime of 15 °C/15 °C. The immature development of female spider mites was prolonged for 7 days at 15 °C/15 °C as compared to 20 °C/5 °C. With a daytime temperature of 20 °C, no differences in lifetime fecundity were observed between a nighttime temperature of 20 and 5 °C for P. persimilis and T. urticae. The two latter species did show a higher lifetime fecundity at 20 °C/5 °C than at 15 °C/15 °C, and their daily fecundity at the alternating regime was about 30 % higher than at the corresponding mean constant temperature. P. persimilis and T. urticae showed no differences in sex ratio between the three temperature regimes, whereas the proportion of N. californicus females at 15 °C/15 °C (54.2 %) was significantly lower than that at 20 °C/5 °C (69.4 %) and 20 °C/20 °C (67.2 %). Intrinsic rates of increase were higher at the alternating temperature than at the corresponding mean constant temperature for both pest and predators. Our results indicate that thermal responses of the studied phytoseiid predators to alternating temperature regimes used in energy saving strategies in glasshouse crops may have consequences for their efficacy in biological control programs.

Acaricides and predatory mites against the begonia mite, Polyphagotarsonemus latus (Acari: Tarsonemidae), on Hedera helix.

In recent years, the begonia mite (Polyphagotarsonemus lotus) has become an important threat to different ornamental cultures in warm greenhouses. At present there are no professional plant protection products registered in Belgium for the control of mites of the Tarsonemidae family. In a screening trial, we evaluated the efficacy of a range of different acaricides: abamectin, milbemectin, pyridaben, spirodiclofen. Based on the results of the screening trial several products were selected for a full efficacy trial following EPPO guidelines. The best control results were obtained with two products from the avermectine group: abamectin and milbemectin. As growers currently have to rely solely on the use of natural enemies there is a strong need for practical evaluation of efficacies of the various predatory mite species (Amblyseius swirskii, A. cucumeris, A. andersoni) used in biological mite control. In a series of experiments, we screened the use of different species of predatory mites. The first efficacy trials on heavily infested plants at different rates of dosage and under different circumstances (temperature, dose rate, application technique) were started in May 2008. In these experiments Amblyseius swirskii showed good efficacy. But temperature was the limiting factor: the predatory mite needed a minimal temperature of 18 degrees C to obtain good results. Further research is necessary to search for predatory mites that can be used in winter conditions (lower temperatures, less light).

Study of use of different types of hydrogen peroxides (2006-2008).

Hydrogen peroxides are commonly used in greenhouses for cleaning purposes and disinfection of irrigation water systems, i.e., to prevent clogging by duckweed (Lemna minor), algae and other (micro)organisms. This use contains a potential risk of involuntary contact to the plants, e.g., to roots through irrigation or to the plant leaves through accidental droplets (spraying mist). To help growers to maximize disinfection with minimal risks, the efficacy and plant safety of a variety of commercial available peroxide formulations were compared, i.e., pure peroxide products, peroxide products with additives: Ag, performic acid, peracetic acid and sorbitol. Starting from pure (clean and without fertilizers) irrigation water the peroxides with Ag-stabilisers were most stable and most effective for algae prevention. In screenings for the curative effect on algae, duckweed and bacteria the best results were obtained with peroxide formulations with performic acid. In plant safety tests on potted Ficus benjamina, sprays and irrigations above the plants gave no toxicity till 500 ppm a.i.; irrigations below the plants didn't show toxicity but the plant growth was reduced with weekly applications of 2000 ppm a.i. On the contrary several applications were risky on herbaceous plants, sometimes even with very low dosages (12.5 ppm peroxide).

Degeneration of sperm reservoir and the loss of mating ability in worker ants.

Workers never mate in the large majority of ants, and they have usually lost the spermatheca, an organ specialized for long-term storage of sperm. Such 'non-sexual' workers are restricted to laying unfertilized eggs that give rise to males, and they cannot compete with the queens for the production of female offspring. In sharp contrast, workers in 200-300 species from phylogenetically basal subfamilies can reproduce sexually ('gamergates') because they retain a functional spermatheca like the queens. Importantly, 'non-sexual' workers in closely related species have a vestigial spermatheca. In this study, we compared the reservoir epithelium of 'sexual' workers to that of congeneric queens and 'non-sexual' workers using 21 species of Amblyoponinae, Ponerinae and Ectatomminae. We show that a pronounced thickening of the epithelium near the opening of the sperm duct is strictly associated with sexual reproduction in both castes. This is unlike 'non-sexual' workers in which this epithelium is always very thin, with few organelles; but all other structures remain intact. We discuss this evolutionary degeneration of the spermatheca and how it relates to behavioural or physiological modifications linked to mating. Our results help understand the loss of sexual reproduction by ant workers, a critical step in the extreme specialization of their phenotype.

The importance of arthropod pests in Belgian pome fruit orchards.

Located in temperate, maritime climate with frequent rainfall, crop protection in Belgian orchards is dominated by fungicides. Though, the importance of arthropod pests should not be underestimated. Pcfruit, the former Research station of Gorsem, has been maintaining a warning system for fruit pests in Belgium since 1944. Therefore, various pests and beneficial's and their life cycle stages have been monitored in Gorsem and in different observation posts across Belgium, being part of a monitoring network. Although up to 3000 arthropod species are present in pome fruit orchards, about 25% can be considered as harmful and another 25% as beneficial. Out of those species, around 100 harmful and 50 beneficial organisms are omnipresent. The list of monitored species is extended yearly for upcoming or difficult to control organisms. Integrated pest management was introduced in the eighties, with the accent on using selective pesticides and saving beneficial organisms. A shift in pesticide use affected the importance of secondary pests, together with recent exceptional climatic conditions. Following many years of monitoring insects and mites and editing warning bulletins in our station, a ranking of the economical importance of different pest species is presented.

Efficacy results of insecticides against Cydia pomonella, the codling moth, in Belgium during the last decade (1998-2007).

The codling moth is an economically important top fruit pest, with its major flight from May till August. We give an overview of ten years of results of efficacy field trials against the codling moth Cydia pomonella, with a number of commonly used products (diflubenzuron, fenoxycarb, flufenoxuron, indoxacarb, granulosis virus, methoxyfenozide and tebufenozide). All trials were executed according to EPPO guidelines. Trials were performed in the Belgian fruit growing area nearby Sint-Truiden, on apple or pear. Each product was sprayed repetitively at registered dose rates to cover the whole flight period of the codling moth, as monitored with pheromone traps. Applications were repeated at strict intervals (7, 14, 21 or 28 days). We show that the different active ingredients had diverse mean efficacies, ranging from 49% to 98%. In general low mean efficacies could be attributed to high variability between trials. Maximal efficacies were indeed high (80%-100%) for all except one "product-interval" combinations, whereas minimal efficacies ranged from 0% to 92%. Belgian fruit growers can thus choose from a large set of active ingredients for resistance management, but have to take variability in efficacy into account. The best and most constant efficacies were reached with Cascade 100 DC (flufenoxuron) at a dose rate of 44 to 50 g a.i./ha leaf wall area and with an application interval of 14 or 21 days.

Sublethal effects of crop protection on honey bee pollination: foraging behaviour and flower visits.

Crop protection strategies essential for pest and disease control can pose risk to pollinators. Fruits cannot be grown commercially without the use of crop protection agents, either from organic or chemical origin. The use of products with toxic effects is banned during flowering, and precise pre-flowering intervals have to be respected in Good Agricultural Practice. Bee pollination is essential for fruit crops to guarantee maximal fruit quality (shape and size) and quantity (yield weight). Fruit growers in Belgium depend mostly on non-commercial beekeepers to provide sufficient colonies for adequate pollination. Under optimal circumstances, beekeepers and fruit growers have mutual benefits from this cooperation as both honey and fruit yield increase. In those European countries with a monitoring scheme, acute bee poisoning incidents have decreased considerably and hardly cause problems at present. In recent years, some concerns arose around sublethal effects (i.e., behavioural changes) of chemical crop protection on bees, especially with regard to increased winter mortality. Even though short-term effects can indeed be induced in individually exposed bees, studies that exposed complete colonies did not reveal any long-term consequences at colony level. However, from the fruit growers' viewpoint, potential short-term effects on foraging behaviour are relevant as they can bear on pollination efficacy.

Queen-worker differences in spermatheca reservoir of phylogenetically basal ants.

Ant queens mate when young and store sperm in their spermatheca to fertilize eggs for several years until their death. In contrast, workers in most species never mate. We have compared the histological organization of spermathecae in 25 poneromorph species exhibiting various degrees of queen-worker dimorphism. The spermathecae of both castes in all species are similar in having a reservoir connected by a sperm duct to the ovary, and a paired gland opening into this duct. The reservoir of queens typically has a columnar epithelium in the hilar region (near the opening of the sperm duct), whereas the epithelium in the distal region is cuboidal. Abundant mitochondria together with apical microvilli and basal invaginations indicate an osmoregulatory function. In contrast, the reservoir epithelium of workers is flattened throughout and lacks these transport characteristics. This single difference shows the importance of a columnar epithelium in the reservoir for sperm storage. However, our data have not revealed inter-specific variations in the development of the hilar region linked with higher fecundity. We have found no consistent differences in associated structures, such as the spermatheca gland or sperm ducts, or in the musculature between queens and workers.

The energetic cost of reproductive conflicts in the ant Pachycondyla obscuricornis.

In a variety of social animals, individuals can secure reproductive rights through aggressive dominance. Direct individual benefits of aggression are widely recognized, but underlying costs affecting group productivity, and thus indirect benefits, are less clear. Costs of aggressive regulation of reproduction are especially important in small social insect colonies, where individual workers could potentially dominate male production. We estimated the energetic costs associated with the regulation of worker reproduction in the ponerine ant Pachycondyla obscuricornis, using the total CO2 emission of a colony as a measure. The level of CO2 emission of 12 experimental colonies varied significantly during five periods with varying levels of aggression and egg-laying. Overall, CO2 emission increased with the degree of fighting in a colony, but was not associated with differences in egg-laying. Aggressive regulation of reproduction and the formation of a dominance hierarchy thus pose an energetic cost to the colony. Furthermore, workers reduce their work-activities immediately after experimental orphaning, giving a further cost to the colony. These costs might influence the outcome of conflicts over male production in ants. This paper presents the first quantification of energetic costs of aggressive behavior regulating reproduction in ants.

Sumo wrestling in ants: major workers fight over male production in Acanthomyrmex ferox.

In the myrmicine ant Acanthomyrmex ferox, major workers have the same number of ovarioles as queens, thrice that of minor workers, making them well suited for egg-laying. In the queen's presence, infrequent aggression allows ranking of majors but they lay only unviable trophic eggs. Major workers engage each other, but not the minors, in antennal boxing and spectacular shaking contests, a novel interaction in ants. The absence of reversals allows a clear ranking of major workers. After queen removal, aggression becomes very frequent, but previous ranking is maintained. All majors start laying reproductive eggs although they show a skew in ovary development according to ranking. The dominant major, however, actively patrols the egg-pile and cannibalises eggs laid by subordinates.

The emergence of collective foraging in the arboreal Gnamptogenys menadensis (Hymenoptera: Formicidae).

Gnamptogenys menadensis is an arboreal nester that forages opportunistically almost exclusively on vegetation, sometimes recruiting others to participate in prey retrieval. The three-dimensional characteristics of vegetation suggest that functions describing recruitment decision thresholds or the pattern of recruitment in arboreal species may differ from those predicted by optimal foraging theory. To examine the effects of prey abundance and distance on the recruitment dynamics of G. menadensis, we baited nests with one termite, five termites or a number of termites between 20 and 40 either near to or far from the entrance and observed the ensuing behaviors. G. menadensis recruited others when encountering multiple termites regardless of the termite pile's distance from the nest, although a few individuals remained at the site and defended the resource. The pattern of arrivals at the site indicates that the majority and sometimes all arrivals were recruited from the branch trails. In combination, these results suggest that the architecture of the foraging habitat, which limits available return routes to the nest and thus increases encounter probabilities with potential recruits, shaped the process of information transfer and generated a collective pattern of foraging and prey retrieval.

Comparative survey of abdominal gland secretions of the ant subfamily Ponerinae.

The chemical contents of three abdominal glands were investigated in representative species of the ponerine ants. The Dufour glands of 14 species show a wide variety of contents. In Mystrium camillae and Proceratium itoi, no volatile substances were found in either the Dufour or venom glands. In Ectatomma sp., Diacamma ceylonense, Diacamma indicum, Pachycondyla obscuricornis, and Pachycondyla striata, volatile chemicals were found in the venom glands as well as in the Dufour glands. Platythyrea punctata was examined, but unusually it does not have a Dufour gland and its venom gland contained no volatile substances. Epoxides were found in ants for the first time in the Dufour glands of Amblyopone reclinata. Venom glands of Pachycondyla tarsata were also found to contain volatile material, including bitter-tasting cyclic dipeptides. In all, 16 species have been added to the list of those examined. All of the 27 known analyses of Dufour glands, 21 analyses of venom glands, and 4 of pygidial glands of workers of ponerine ant species have been brought together in order to seek some pattern in the type of glandular contents. Although the great majority of species produce hydrocarbons in their worker Dufour glands, and some have terpenes, there is no observable pattern for this gland on a tribe or genus level. Volatile compounds have been found in the venom glands of some species of the tribe Ponerini only. The information on pygidial glands is still too fragmentary for any conclusions.