Neoseiulus paspalivorus,a predator from coconut,as a candidate for controlling dry bulb mites infesting stored tulip bulbs

The dry bulb mite, Aceria tulipae, is the most important pest of stored tulip bulbs in The Netherlands. This tiny, eriophyoid mite hides in the narrow space between scales in the interior of the bulb. To achieve biological control of this hidden pest, candidate predators small enough to move in between the bulb scales are required. Earlier experiments have shown this potential for the phytoseiid mite, Neoseiulus cucumeris, but only after the bulbs were exposed to ethylene, a plant hormone that causes a slight increase in the distance between tulip bulb scales, just sufficient to allow this predator to reach the interior part of the bulb. Applying ethylene, however, is not an option in practice because it causes malformation of tulip flowers. In fact, to prevent this cosmetic damage, bulb growers ventilate rooms where tulip bulbs are stored, thereby removing ethylene produced by the bulbs (e.g. in response to mite or fungus infestation). Recently, studies on the role of predatory mites in controlling another eriophyoid mite on coconuts led to the discovery of an exceptionally small phytoseiid mite, Neoseiulus paspalivorus. This predator is able to move under the perianth of coconuts where coconut mites feed on meristematic tissue of the fruit. This discovery prompted us to test N. paspalivorus for its ability to control A. tulipae on tulip bulbs under storage conditions (ventilated rooms with bulbs in open boxes; 23 °C; storage period June–October). Using destructive sampling we monitored predator and prey populations in two series of replicated experiments, one at a high initial level of dry bulb mite infestation, late in the storage period, and another at a low initial dry bulb mite infestation, halfway the storage period. The first and the second series involved treatment with N. paspalivorus and a control experiment, but the second series had an additional treatment in which the predator N. cucumeris was released. Taking the two series of experiments together we found that N. paspalivorus controlled the populations of dry bulb mites both on the outer scale of the bulbs as well as in the interior part of the bulbs, whereas N. cucumeris significantly reduced the population of dry bulb mites on the outer scale, but not in the interior part of the bulb. Moreover, N. paspalivorus was found predominantly inside the bulb, whereas N. cucumeris was only found on the outer scale, thereby confirming our hypothesis that the small size of N. paspalivorus facilitates access to the interior of the bulbs. We argue that N. paspalivorus is a promising candidate for the biological control of dry bulb mites on tulip bulbs under storage conditions in the Netherlands.     

Anthocorid predators learn to associate herbivore-induced plant volatiles with presence or absence of prey

We investigated how the plant‐inhabiting, anthocorid predator, Anthocoris nemoralis, copes with variation in prey, host plant and associated herbivore‐induced plant volatiles and in particular whether the preference for these plant odours is innate or acquired. We found a marked difference between the olfactory response of orchard‐caught predators and that of their first generation reared on flour moth eggs in the laboratory, i.e. under conditions free of herbivory‐induced volatiles. Whereas the orchard‐caught predators preferred odour from psyllid‐infested pear leaves, when offered against clean air in a Y‐tube olfactometer, the laboratory‐reared first generation of (naive) predators did not. The same difference was found when a single component (methyl salicylate) of the herbivore‐induced plant volatiles was offered against clean air. After experiencing methyl salicylate with prey, however, the laboratory‐reared predators showed a pronounced preference for this volatile. This acquired preference did not depend on whether the volatile had been experienced in the juvenile period or in the adult phase, but it did depend on whether it had been offered in presence or absence of prey. In the first case, they were attracted to the plant volatile in subsequent olfactometer experiments, but when the volatile had been offered during a period of prey deprivation, the predators were not attracted. We conclude that associative learning is the most likely mechanism underlying acquired odour preference.     

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 ).     

Alternative phenotypes of male mating behaviour in the two-spotted spider mite

Severe intraspecific competition for mates selects for aggressive individuals but may also lead to the evolution of alternative phenotypes that do not act aggressively, yet manage to acquire matings. The two-spotted spider mite, Tetranychus urticae, shows male mate-guarding behaviour and male–male combat for available females. This may provide opportunity for weaker males to avoid fighting by adopting alternative mating behaviour such as sneaker or satellite tactics as observed in other animals. We investigated male precopulatory behaviour in the two-spotted spider mite by means of video-techniques and found three types of male mating behaviour: territorial, sneaker and opportunistic. Territorial and sneaker males associate with female teleiochrysales and spend much time guarding them. Territorial males are easily disturbed by rival males and engage themselves in fights with them. However, sneaker males are not at all disturbed by rival males, never engage in fights and, strikingly, never face attack by territorial males. Opportunistic males wander around in search of females that are in the teleiochrysalis stage but very close to or at emergence. To quickly classify any given mate-guarding male as territorial or sneaker we developed a method based on the instantaneous response of males to disturbance by a live male mounted on top of a brush. We tested this method against the response of the same males to natural disturbance by two or three other males. Because this method proved to be successful, we used it to collect territorial and sneaker males, and subjected them to morphological analysis to assess whether the various behavioural phenotypes are associated with different morphological characters. However, we found no statistical differences between territorial and sneaker males, concerning the length of the first legs, the stylets, the pedipalps or the body.     

Modelling Fungal (Neozygites cf. Floridana) Epizootics in Local Populations of Cassava Green Mites (Mononychellus Tanajoa)

The fungus, Neozygitis cf. floridana is parasitic on the cassava green mite, Mononychellus tanajoa (Bondar) (Acari: Tetranychidae) in South America and may be considered for classical biological control of cassava green mites in Africa, where cassava is an important subsistence crop, cassava green mites are an imported pest and specific natural enemies are lacking. Spider mites generally have a viscous structure of local populations, a trait that would normally hamper the spread of a fungus that is transmitted by the contact of susceptible hosts with the halo of capilliconidia surrounding an infectious host. However, if infected mites search and settle to produce capilliconidia on sites where they are surrounded by susceptible mites before becoming infectious, then the conditions for maximal transmission in a viscous host population are met. Because the ratio between spider mites and the leaf area they occupy is constant, parasite-induced host searching behaviour leads to a constant per capita transmission rate. Hence, the transmission rate only depends on the number of infectious hosts. These assumptions on parasite-induced host search and constant host density lead to a simple, analytically tractable model that can be used to estimate the maximal capacity of the fungus to decimate local populations of the cassava green mite. By estimating the parameters of this model (host density, per capita transmission rate and duration of infected and infectious state) it was shown that the fungal pathogen can reduce the population growth of M. tanajoa, but cannot drive local mite populations to extinction. Only when the initial ratio of infectious to susceptible mites exceeds unity or the effective growth rate of the mite population is sufficiently reduced by other factors than the fungus (e.g. lower food quality of the host plant, dislodgement and death by rain and wind and predation), will the fungal pathogen be capable of decimating the cassava green mite population. Under realistic field conditions, where all of these growth-reducing factors are likely to operate, there may well be room for effective control by the parasitic fungus. This revised version was published online in November 2006 with corrections to the Cover Date.     

Does prey preference change as a result of prey species being presented together? Analysis of prey selection by the predatory mite Typhlodromus pyri (Acarina: Phytoseiidae)

Summary Prey-selection behaviour of the phytoseiid mite Typhlodromus pyri Scheuten was analysed with a Markovtype model of feeding-state dynamics and feeding-state dependent searching behaviour (Sabelis 1981, 1986, 1989; Metz and Van Batenburg 1985a, b). All behavioural characteristics of the predator which are independent of the feeding state were represented by one parameter. The remaining feeding-state dependent characteristics were represented by a function of the feeding state, with one parameter. The best parameter values to describe a predator-prey interaction were determined by fitting the model to the predation rates in monocultures. Under the assumption that the parameter values are not dependent on the composition of prey species supply, the diet of the predators in mixed cultures was predicted from parameters estimated in monoculture experiments.Two prey types, apple rust mite (Aculus schlechtendali (Nalepa)) adults and European red spider mite (Panonychus ulmi (Koch)) larvae were studied. A large discrepancy was observed between calculated and experimentally determined predation rates of T. pyri in mixed cultures: the predators actually killed 3–7 times more P. ulmi larvae than was predicted by the model.The large difference between observed and predicted predation rates in mixed cultures cannot be explained by changes in the behaviour of the prey species as a result of being together. Therefore, it seems likely that the prey selection behaviour of the predator was different when prey species were presented together than when presented singly. Apparently the predatory mite T. pyri prefers P. ulmi to S. schlechtendali.     

Karyotype displacement in a laboratory population of the two spotted spider mite Tetranychus urticae (Koch): Experiments and computer simulations

Three different strains, homozygous for a radiation induced structural chromosome mutation (T), exhibiting negative hererosis, were tested for their ability to displace the standard (wild-type) karyotype from experimental populations. The experimental populations were initiated by mixing fertilized females of both a T strain and the standard strain at different ratios. Two of the T strains showed the ability to displace the standard karyotype if the initial frequency of the T karyotype was at least 0.65. The additional release of T males into the experimental population accelerated considerably the process of displacement of the standard karyotype, especially if the initial T karyotype frequency was 0.65.A computer model for simulating the process of population displacement in Tetranychus urticae was developed. The model accounts for variation in developmental time and for the age dependency of variables related to fitness. The simulations showed a good correlation with the experimental results. A system analysis on the sensitivity of the model output to varying different population parameters demonstrated that especially the relative number of males produced by a T strain and the female developmental rate were of significant importance to the population displacement ability of a T strain. The negative influence of genetic markers on general fitness and various aspects of practical application of the method are discussed.     

Limits to runaway sexual selection: The wallflower paradox

In his mathematical treatment of Fisher's ideas on sexual selection (so-called runaway selection) Lande (1981) predicted that males may evolve increasingly elaborate sexual characters despite opposing viability selection as a consequence of the associated costs. Lande thereby assumed that female mate preferences are not subject to selection since (1) females are all inseminated and (2) the quantity and quality of their offspring are independent of the female's mate preferences. Kirkpatrick (1985) removed the latter assumption and investigated the consequences for the mean phenotype with respect to both female and male traits. He also explored the dynamics of the (co)-variance matrix by numerical methods. In this paper we consider a simpler model with just two multi-allelic loci. This enables us to derive explicit expressions for (co)-variances under steady state conditions. Rather than assume natural selection through differential fertility (as in Kirkpatrick, 1985), we take sexual selection on females into account by modelling the preference-dependent risk that females remain unmated. We argue that this wallflower effect is a realistic feature of any mating system, since it merely depends on the existence of (1) variation in mating preferences and (2) a finite mating season. Our approach provided an insight into the dynamic behaviour of the means of the phenotypes. This is because the dynamics of the means depend on the steady state (co)-variance matrix. Thus, an insight into the former requires explicit expressions for the latter. Whereas Lande and Kirkpatrick predicted runaway processes, despite opposing viability selection, our model predicts a globally stable steady state, i.e. no runaway, even without opposing viability selection (under the assumption of an asymptotically stable steady state of the (co)-variances. Admittedly, we have no analytic proof of this stability but only support for it, based on simulations.) The absence of the runaway processes in our model is caused by the wallflower effect, since it imposes constraints on the steady state of the (co)-variance matrix. When mutational input applies to female traits but not to male traits, explicit expressions for the (co)-variances under steady state conditions can be derived, and these show that: (1) both the genetic covariance and the variance of male traits are equal to zero, but (2) the variance of the female trait exceeds to zero. Should there be mutational input influencing the male trait, then these results would suggest that the male-to-female ratio of variances is much smaller than unity. This prediction is of tremendous importance for speciation through founding events.