Variability in susceptibility among cucumber and tomato strains of Tetranychus urticae Koch to 2-tridecanone from tomato trichomes: effects of host plant shift

The effects of secondary plant compounds on different host races/strains of a herbivorous arthropod are not easy to interpret based on dose–response tests alone. This difficulty arises because the responses in a population to a given dose are dictated not only by genetic factors, but also by factors such as feeding history, age and plant. To discriminate between these possibilities different strains of a herbivorous mite (Tetranychus urticae Koch) were exposed to one relevant toxin from glandular hairs on tomato (the methyl ketone, 2-tridecanone) and these strains had a known feeding history on either or both of two host plants (tomato and cucumber). It was hypothesized that tomato is a relatively hostile host plant to spider mites partly due to methyl ketones from glandular hairs and that consequently there will be stronger selection on the tomato strains than on the cucumber strains. However, the expected differences between the tomato and cucumber strains did not show up; three spider mite strains collected from tomato and two strains collected from cucumber appeared to be equally susceptible to 2-tridecanone. This unexpected result cannot result from selection for resistance to 2-tridecanone, but it may be the consequence of cross-resistance to other ketones in cucumber or the bio-accumulation (i.e. metabolic load) of 2-tridecanone prior to the toxicity test. To test this hypothesis, one of the tomato strains was released on cucumber for different time intervals and one of the cucumber strains on tomato. It was found that the resistance of the tomato strain to 2-tridecanone increased 6 months after transfer to cucumber. This increased resistance cannot be the result of selection because 2-tridecanone is absent from cucumber. Hence, it may be due either to selection for resistance to another ketone in cucumber, possibly leading to cross-resistance, or to the absence of 2-tridecanone bioaccumulation on cucumber. Transfer of the cucumber strain from cucumber to tomato also increased the resistance to 2-tridecanone. As this was accompanied by high mortality directly after the transfer, selection for resistance may have played a role. Alternatively, the increased resistance may be due to induction of resistance to secondary plant compounds of tomato, including 2-tridecanone. In conclusion, experiments on host plant transfer show that the tomato strain and the cucumber strain are not equal in their resistance to 2-tridecanone.     

Plant structural changes due to herbivory: Do changes in <Emphasis Type="Italic">Aceria</Emphasis>-infested coconut fruits allow predatory mites to move under the perianth?

Being minute in size, eriophyoid mites can reach places that are small enough to be inaccessible to their predators. The coconut mite, Aceria guerreronis, is a typical example; it finds partial refuge under the perianth of the coconut fruit. However, some predators can move under the perianth of the coconut fruits and attack the coconut mite. In Sri Lanka, the phytoseiid mite Neoseiulus baraki, is the most common predatory mite found in association with the coconut mite. The cross-diameter of this predatory mite is c. 3 times larger than that of the coconut mite. Nevertheless, taking this predator’s flat body and elongated idiosoma into account, it is—relative to many other phytoseiid mites—better able to reach the narrow space under the perianth of infested coconut fruits. On uninfested coconut fruits, however, they are hardly ever observed under the perianth. Prompted by earlier work on the accessibility of tulip bulbs to another eriophyoid mite and its predators, we hypothesized that the structure of the coconut fruit perianth is changed in response to damage by eriophyoid mites and as a result predatory mites are better able to enter under the perianth of infested coconut fruits. This was tested in an experiment where we measured the gap between the rim of the perianth and the coconut fruit surface in three cultivars (‘Sri Lanka Tall’, ‘Sri Lanka Dwarf Green’ and ‘Sri Lanka Dwarf Green × Sri Lanka Tall’ hybrid) that are cultivated extensively in Sri Lanka. It was found that the perianth-fruit gap in uninfested coconut fruits was significantly different between cultivars: the cultivar ‘Sri Lanka Dwarf Green’ with its smaller and more elongated coconut fruits had a larger perianth-fruit gap. In the uninfested coconut fruits this gap was large enough for the coconut mite to creep under the perianth, yet too small for its predator N. baraki. However, when the coconut fruits were infested by coconut mites, the perianth-rim-fruit gap was not different among cultivars and had increased to such an extent that the space under the perianth became accessible to the predatory mites.     

Predators induce egg retention in prey

To prevent predation on their eggs, prey often avoid patches occupied by predators. As a result, they need to delay oviposition until they reach predator-free patches. Because many species allocate energy to egg production in a continuous fashion, it is not clear what kind of mechanism prey use to delay oviposition. We used females of the phytoseiid mite Neoseiulus cucumeris to study these mechanisms. Females were placed in patches with pollen, a food source they use for egg production, and they were exposed to another phytoseiid mite, Iphiseius degenerans, which is an intraguild predator of N. cucumeris juveniles. We found that the oviposition of N. cucumeris females on patches with the predator was lower than on patches without the predator. Cues left by the intraguild predator were not sufficient to elicit such behaviour. Females of N. cucumeris reduced oviposition when exposed to the predator by retaining the egg inside their body, resulting in a lower developmental rate once these eggs were laid. Hence, females are capable of retaining eggs, but the development of these eggs continues inside the mother’s body. In this way, females gain some time to search for less risky oviposition sites.     

How predator functional responses and Allee effects in prey affect the paradox of enrichment and population collapses

In Rosenzweig-MacArthur models of predator-prey dynamics, Allee effects in prey usually destabilize interior equilibria and can suppress or enhance limit cycles typical of the paradox of enrichment. We re-evaluate these conclusions through a complete classification of a wide range of Allee effects in prey and predator's functional response shapes. We show that abrupt and deterministic system collapses not preceded by fluctuating predator-prey dynamics occur for sufficiently steep type III functional responses and strong Allee effects (with unstable lower equilibrium in prey dynamics). This phenomenon arises as type III functional responses greatly reduce cyclic dynamics and strong Allee effects promote deterministic collapses. These collapses occur with decreasing predator mortality and/or increasing susceptibility of the prey to fall below the threshold Allee density (e.g. due to increased carrying capacity or the Allee threshold itself). On the other hand, weak Allee effects (without unstable equilibrium in prey dynamics) enlarge the range of carrying capacities for which the cycles occur if predators exhibit decelerating functional responses. We discuss the results in the light of conservation strategies, eradication of alien species, and successful introduction of biocontrol agents.     

Diapause incidence in the two-spotted spider mite increases due to predator presence, not due to selective predation

We recently reported evidence for increased diapause incidence in the spider mite Tetranychus urticae in presence of the predatory mite Typhlodromus pyri. This effect may arise from (1) selective predation on non-diapause spider mites, (2) predator-induced diapause in spider mites, or (3) both. Using a different strain of T. urticae, we first recovered increased diapause incidence in association with predators. Then, we tested for selective feeding in two-choice experiments with equal numbers of non-diapause and diapause spider mites. We found that the predatory mite had a significant preference for the latter. This indicates that increased diapause incidence in association with predatory mites is not due to selective predation. Therefore, predator-mediated physiological induction of diapause seems a more likely explanation. The cues leading to induction appear to relate to the predators, not their effects, since predation simulated by spider-mite removal or puncturing did not significantly affect diapause incidence. Why spider mites benefit from this response, remains an open question.This revised version was published online in May 2005 with a corrected cover date.     

Gross morphology of the central nervous system of a phytoseiid mite

The general morphology of the central nervous system is analysed in intact females of the predatory mite, Phytoseiulus persimilis (Acari: Phytoseiidae), using a nucleic acid label (YOYO-1) and confocal laser scanning microscopy. The somata of all cells that comprise the synganglion reside in the cortex. The cortex harbours an estimated total of 10,000 cells. The somata are densely packed in the cortex and cells residing in the inner cortex may only occupy about 1.8 μm. As in all Arachnida, the synganglion is divided in a sub- and a supra-oesophageal nervous mass. Both the cortex and the neuropil appear continuous between these two nervous masses. The sub-oesophageal nervous mass mainly consists of the four paired pedal ganglia that are each associated with a leg. The prominent olfactory lobes are ventrally associated with the first pedal ganglia. A small opisthosomal ganglion occupies the most caudal part of the sub-oesophageal ganglion. The rostral part of the supra-oesophageal nervous mass consists of the paired cheliceral and palpal ganglia. The supra-oesophageal ganglion is the largest ganglion in the supra-oesophageal nervous mass and unlike all other ganglia it is not associated with any of the major nerves. It is therefore more likely involved in secondary information processing.     

Laboratory tests for controlling poultry red mites (Dermanyssus gallinae) with predatory mites in small ‘laying hen’ cages

To assess their potential to control poultry red mites (Dermanyssus gallinae), we tested selected predaceous mites (Androlaelaps casalis and Stratiolaelaps scimitus) that occur naturally in wild bird nests or sometimes spontaneously invade poultry houses. This was done under laboratory conditions in cages, each with 2–3 laying hens, initially 300 poultry red mites and later the release of 1,000 predators. These small-scale tests were designed to prevent mite escape from the cages and they were carried out in three replicates at each of three temperature regimes: 26, 30 (constant day and night) and 33–25?°C (day-night cycle). After 6?weeks total population sizes of poultry red mites and predatory mites were assessed. For the temperature regimes of 26 and 33/25?°C S. scimitus reduced the poultry red mite population relative to the control experiments by a factor 3 and 30, respectively, and A. casalis by a factor of 18 and 55, respectively. At 30?°C the predators had less effect on red mites, with a reduction of 1.3-fold for S. scimitus and 5.6-fold for A. casalis. This possibly reflected hen manure condition or an effect of other invertebrates in the hen feed. Poultry red mite control was not negatively affected by temperatures as high as 33?°C and was always better in trials with A. casalis than in those with S. scimitus. In none of the experiments predators managed to eradicate the population of poultry red mites. This may be due to a prey refuge effect since most predatory mites were found in and around the manure tray at the bottom of the cage, whereas most poultry red mites were found higher up in the cage (i.e. on the walls, the cover, the perch, the nest box and the food box). The efficacy of applying predatory mites in the poultry industry may be promoted by reducing this refuge effect, boosting predatory mite populations using alternative prey and prolonged predator release devices. Biocontrol success, however, will strongly depend on how the poultry is housed in practice (free range, cage or aviary systems) and on which chemicals are applied to disinfect poultry houses and to control other pests.     

Herbivore arthropods benefit from vectoring plant viruses

Plants infected with pathogens often attract the pathogens’ vectors, but it is not clear if this is advantageous to the vectors. We therefore quantified the direct and indirect (through the host plant) effects of a pathogen on its vector. A positive direct effect of the plant‐pathogenic Tomato spotted wilt virus on its thrips vector (Frankliniella occidentalis) was found, but the main effect was indirect; juvenile survival and developmental rate of thrips was lower on pepper plants that were damaged by virus‐free thrips than on unattacked plants, but such negative effects were absent on plants that were damaged and inoculated by infected thrips or were mechanically inoculated with the virus. Hence, potential vectors benefit from attacking plants with virus because virus‐infected plants are of higher quality for the vector's offspring. We propose that plant pathogens in general have evolved mechanisms to overcome plant defences against their vectors, thus promoting pathogen spread.     

Plants protect their roots by alerting the enemies of grubs

Plant roots in the soil are under attack from many soil organisms. Although many ecologists are aware of the presence and importance of natural enemies in the soil that protect the plants from herbivores, the existence and nature of tritrophic interactions are poorly understood. So far, attention has focused on how plants protect their above-ground parts against herbivorous arthropods, either directly or indirectly (i.e. by getting help from the herbivore's enemies). This article is the first in showing that indirect plant defences also operate underground. We show that the roots of a coniferous plant ( Thuja occidentalis ) release chemicals upon attack by weevil larvae ( Otiorhynchus sulcatus ) and that these chemicals thereby attract parasitic nematodes ( Heterorhabditis megidis ).     

Dynamics of refuge use: diurnal, vertical migration by predatory and herbivorous mites within cassava plants

Plants may protect themselves against herbivorous arthropods by providing refuges to predatory arthropods, but they cannot prevent herbivores from taking countermeasures or even from reaping the benefits. To understand whether plants benefit from providing self‐made refuges (so‐called domatia), it is not only necessary to determine the fitness consequences for the plant, but also to assess (1) against which factors the refuge provides protection, (2) why predatory arthropods are more likely to monopolise the refuge, and (3) how herbivorous and predatory arthropods respond to and affect each other in and outside the refuge. In this article, we focus on the last aspect by studying the dynamics of refuge use of a predatory mite (Typhlodromalus aripo) and its consequences for a herbivorous mite (Mononychellus tanajoa) on cassava plants in Benin, West Africa. The refuge, located in‐between the leaf primordia of the cassava apex, is thought to provide protection against abiotic factors and/or intraguild predators. To test whether the predator waits for prey in the apex or comes out, we sampled predator‐prey distributions on leaves and in the apex at 4 hour‐intervals over a period of 24 hours. The predatory mites showed pronounced diurnal changes in within‐plant distribution. They were in the apices during the day, moved to the young leaves during night and returned to the apices the next morning. Nocturnal foraging bouts were more frequent when there were more herbivorous mites on the leaves near the apex. However, the foraging predators elicited an avoidance response by mobile stages of their prey, since these were more abundant on the first 20 leaves below the apex during late afternoon, than on the same leaves during night. These field observations on cassava plants show that (1) during daytime predatory mites monopolise the apical domatia, (2) they forage on young leaves during night and (3) elicit avoidance by within‐plant, vertical migration of mobile stages of the herbivorous mites. We hypothesize that cassava plants benefit from apical domatia by acquiring protection for their photosynthetically most active, young parts, because predatory mites (1) protect primordial leaves in the apex, (2) reduce the densities of herbivorous mites on young leaves, and (3) cause herbivorous mites to move down to less profitable older leaves.