Vertical Dispersal of Steinernema scapterisci

When infective juveniles ofSteinernema scapterisci Nguyen &Smart were released on the soil surface in the field and in the laboratory, some of them moved downward through the soil at least 10 cm in 5 days and infected and killed mole crickets. When released 2 cm below the soil surface, most of the juveniles moved into the upper 2 cm layer of soil, but some moved downward 10 cm. When placed at the center of a 16-cm soil column, infective juveniles moved in both directions with three times more moving downward than upward. Infective juveniles were more efficient in killing mole crickets in the field than in the laboratory.     

Natural selection in the laboratory for a change in resistance byDrosophila melanogaster to the parasitoid waspAsobara tabida

The selection response of the polymorphic hostD. melanogaster (Meigen) to the braconid waspA. tabida (Nees) is addressed. Cages of flies with and without wasps were initiated with a population ofD. melanogaster that exhibited variation both in larval foraging behavior and in encapsulation ability. Encapsulation ability was measured as the proportion of parasitized larvae that produce a hardened capsule which encapsulates the wasp egg and ultimately kills the wasp larva. We determined whether the host population changed its encapsulation ability and/or its foraging behavior in response to the wasp. Both species were collected from a local orchard whereA. tabida is the only wasp known to parasitizeD. melanogaster larvae. The naturally occurring genetic polymorphism for rover and sitter larval foraging behavior inD. melanogaster is also found in this field population.A. tabida's vibrotactic search behavior enables it to detect rover more frequently than sitter larvae. Rover larvae move significantly more while feeding than do sitter larvae. In this field population, rover larvae also show higher encapsulation abilities than do sitter larvae. Six cage populations, three without wasps and three with wasps, each containing an equal mixture of rover and sitter flies, were established in the laboratory and maintained for 19 fly generations. Selection pressure in the laboratory was similar to that found in the field population from which the flies and wasps were derived. We found that larvae from cages with wasps developed a significantly higher frequency of encapsulation than those reared without wasps. We were, however, unable to detect a change in larval movement (rover or sitter behavior) in larvae from cages subject to selection from wasps compared to larvae from cages containing no wasps. This may have resulted from a balance between two selective forces, selection against rovers by the wasps' use of vibrotaxis, and selection for rovers resulting from their increased encapsulation abilities     

Danger detection and escape behaviour in wood crickets

The wind-sensitive cercal system of Orthopteroid insects that mediates the detection of the approach of a predator is a very sensitive sensory system. It has been intensively analysed from a behavioural and neurobiological point of view, and constitutes a classical model system in neuroethology. The escape behaviour is triggered in orthopteroids by the detection of air-currents produced by approaching objects, allowing these insects to keep away from potential dangers. Nevertheless, escape behaviour has not been studied in terms of success. Moreover, an attacking predator is more than “air movement”, it is also a visible moving entity. The sensory basis of predator detection is thus probably more complex than the perception of air movement by the cerci. We have used a piston mimicking an attacking running predator for a quantitative evaluation of the escape behaviour of wood crickets Nemobius sylvestris. The movement of the piston not only generates air movement, but it can be seen by the insect and can touch it as a natural predator. This procedure allowed us to study the escape behaviour in terms of detection and also in terms of success. Our results showed that 5-52% of crickets that detected the piston thrust were indeed touched. Crickets escaped to stimulation from behind better than to a stimulation from the front, even though they detected the approaching object similarly in both cases. After cerci ablation, 48% crickets were still able to detect a piston approaching from behind (compared with 79% of detection in intact insects) and 24% crickets escaped successfully (compared with 62% in the case of intact insects). So, cerci play a major role in the detection of an approaching object but other mechanoreceptors or sensory modalities are implicated in this detection. It is not possible to assure that other sensory modalities participate (in the case of intact animals) in the behaviour; rather, than in the absence of cerci other sensory modalities can partially mediate the behaviour. Nevertheless, neither antennae nor eyes seem to be used for detecting approaching objects, as their inactivation did not reduce their detection and escape abilities in the presence of cerci.     

Variation between laboratory populations in the performance of the parasitoid Encarsia formosa on two host species, Bemisia tabaci and Trialeurodes vaporariorum

We tested the hypothesis that populations of the parthenogenetic parasitic wasp Encarsia formosa Gahan (Hymenoptera: Aphelinidae) differed in their ability to use two different host species, Bemisia tabaci Gennadius (Homoptera: Aleyrodidae) and Trialeurodes vaporariorum Westwood (Homoptera: Aleyrodidae). Of the three wasp populations tested, two populations had been reared for many generations on B. tabaci and one population had been reared for many years on T. vaporariorum. Performance was measured by the number of whitefly nymphs that were successfully parasitized by individual wasps, and performance on either host was measured in separate experiments. There was variation between wasp populations in their performance on the host B. tabaci, with one wasp population reared for many years on this host performing considerably better than the other two populations. There were no significant differences between populations in their use of the preferred host, T. vaporariorum. The experiments were conducted in such a way that we could distinguish heritable differences between populations from environmentally-induced conditioning differences due to the immediate host from which an individual wasp enclosed. In either experiment there were no significant effects of conditioning, although there was a trend within each population for wasps conditioned on T. vaporariorum to have higher performance than those conditioned on B. tabaci. Thirdly, we conducted a selection experiment, initiated with wasps from a single population historically reared on T. vaporariorum, to measure the effect of laboratory rearing on different hosts for 17 generations. We did not see any difference in the performance of wasps on B. tabaci after this period of rearing on either of the two hosts. In summary, populations of E. formosa do differ in their relative performance on B. tabaci. The one population that was tested further did not show any response to selection by rearing, but the ability to respond to selection on performance may not be equal for all populations. The possibility that wasp populations have differential performance on particular hosts may affect the use of this species as a biological control agent.     

Catching Ariadne by her thread: how a parasitoid exploits the herbivore's marking trails to locate its host

Chemical signals that can be associated with the presence of a host insect often work as arrestants in close range host location by parasitoids, leading to longer searching times on patches where such signals are present. Our current view of parasitoid host location is that by prolonging the search times in patches, randomly searching parasitoids enhance their chance of detecting host insects. However, prolonged search times are not necessarily the only modification in parasitoid behaviour. In this study, we examine the exploitation of host-fruit marking pheromone of rose-hip flies, Rhagoletis basiolaOsten-Sacken (Diptera: Tephritidae) by the specialized egg-larval parasitoid Halticoptera rosae Burks (Hymenoptera: Pteromalidae). We provide evidence that the instantaneous probability that a host egg will be located by a searching parasitoid wasp differs markedly between pheromone-marked and unmarked fruits. The arresting response to the marking pheromone, i.e., the prolonged time a wasp is willing to search on marked fruits, can only account for a small fraction of the difference in successful host location on marked and unmarked fruits. We further demonstrate that the time wasps require to locate the host egg is independent of the size of the rose-hip harbouring the fly egg, and thus is independent of the area the wasp potentially has to search. A comparison of our findings with results of different search algorithms for parasitoid wasps suggests that wasps use the fly's pheromone marking trail as a guide way to the fly's oviposition site and thus the host egg.     

Spiders of the genus Cupiennius Simon 1891 (Araneae,Ctenidae)

Summary Cupiennius salei (Ctenidae) is a tropical wandering spider which lives in close association with a particular type of plant (see companion paper). These plants are the channels through which the spiders receive and emit various types of vibrations. We measured the vibrations the spiders are typically exposed to when they sit on their dwelling plants (banana plant, bromeliad) in their natural biotope in Central America. In addition a laboratory analysis was carried out to get an approximate idea of the complex vibration-propagating properties of the dwelling plants, taking a banana plant as an example. (1) Types of vibrations (Figs. 1–4). Despite variability in detail there are characteristic differences in spectral composition between the vibrations of various abiotic and biotic origins: (a) Vibrations due to wind are very low frequency phenomena. Their frequency spectra are conspicuously narrow with prominent peaks close to or, more often, below 10 Hz. Vibrations due to raindrops show maximal acceleration values at ca. 1000 Hz. Their frequency band at-20 dB extends up to ca. 250 Hz where-as that of the vibrations due to wind extends to only ca. 50 Hz. (b) The frequency spectra of prey vibrations such as those generated by a running cockroach are typically broad-banded and contain high frequencies; they have largest peaks mostly between ca. 400 and 900 Hz. Their-20 dB frequency bands usually extend from a few Hz to ca. 900 Hz. Some potential prey animals such as grass-hoppers seem to be vibrocryptic; they walk by the spider as if unnoticed. Their cautious gait leads to only weak vibrations at very low frequencies resembling the background noise due to wind. Courtship signals are composed maily of low frequencies, intermediate between background noise and prey vibrations (male: prominent peaks at ca. 75 Hz and ca. 115 Hz; female: dominant frequencies between ca. 20 Hz and ca. 50 Hz). The male signal is composed of syllables and differs from all other vibrations studied here by being temporally highly ordered. A comparison with previous electrophysiological studies suggests that the high pass characteristics of the vibration receptors enhance the signal-to-(abiotic)-noise ratio and that the vibration-sensitive interneurons so far examined and found to have band pass characteristics are tuned to the frequencies found in the vibrations of biotic origin. (2) Signal propagation (Fig. 5). In terms of frequency-dependent attenuation of vibrations the banana plant is well suited for transmitting the above signals. Average attenuation values are ca. 0.35 dB/cm. Together with known data on vibration receptor sensitivity this explains the range of courtship signals of more than 1 m observed in behavioral studies. Attenuation in the plant is neither a monotonic function of frequency nor of distance from the signal source.     

Host examination walk and oviposition site selection ofTrichogramma minutum: Studies on spherical hosts

The host examination behavior of the parasitoid wasp Trichogramma minutumon spherical Manduca sextahosts is described. The mean walking speed during host examination was 0.64 ± 0.03 mm s ^–1 and is independent of wasp size. The paths taken by the wasps are not evenly distributed over the host surface; the majority of time is spent below 45 ° latitude. The distribution of oviposition sites is also nonrandom with respect to latitude and is not influenced by phototaxic and geotaxic responses. The initial transit made by the wasps over the host surface is a straight path which frequently passes across the highest point on the host. The length and duration of the initial transit are independent of wasp body length. However, stride length and stepping rates are proportional to wasp body length, and small wasps take more steps to complete their initial transit. The roles of the examination walk in host recognition and host volume measurement by Trichogrammaare also discussed.     

Chemosensory attraction of fig wasps to substances produced by receptive figs

In the mutualism between figs (Ficus spp., Moraceae) and their species-specific fig wasp pollinators (Hymenoptera: Agaonidae), location of a receptive host tree by the adult insect is a critical step. The adult female wasp lives only a few days, and must usually fly to a different tree than her natal tree to locate receptive figs. Trees in receptive phase often occur at very low densities. Reproductive success of both fig and wasp depends on trasmission of a very strong signal by the plant. Some evidence exists for the role of olfaction in location of receptive hosts by fig wasps, but very little work has been done on the chemical ecology of host location and host specificity. Here the first experimental evidence is presented for long-distance olfactory attraction of wasps by volatile substances produced by receptive figs, and for short-distance or contact chemostimulation by host volatiles that elicit entry of the wasp into the fig. In studies usingFicus carica L., pentane extracts of receptive-phase figs attract the pollinatorBlastophaga psenes L. from distances of at least 5 m in the field. Short-distance chemostimulation was demonstrated in laboratory bioassays. Pentane extracts of receptive figs, when painted onto the ostiole of non-receptive figs, elicit entry of pollinator wasps. Figs emit volatile compounds attractive to pollinating wasps only during the period of receptivity; pentane extracts of non-receptive figs are not attractive. A simple reliable procedure is described to compare the attractivity of different types of extracts (total, internal, and external extracts) and of different fractions, in the first step towards identifying attractant substances.     

Homing in the field cricket,Gryllus campestris

Homing ability of the cricket Gryllus campestriswas experimentally studied in the field after passive and active displacements. Whatever the weather conditions and the nature of the displacement, crickets (18/18) home directly when they are located between 5 and 15 cm from their burrow by performing real-time orientation. When moved by the experimenter between 15 and 55 cm away from home, some crickets (26/43) can retrieve their burrow by systematic searching only after winding inward trips but they get lost (0/18) beyond 55 cm. After self-generated movements between 5 and 85 cm away from home, crickets (33/35) return to their burrow under blue sky, but they cannot home as well under overcast sky (22/34 from 5 to 65 cm and 0/5 beyond). The volatile memory orientation system allowing path integration is based on the polarized skylight of the sun.     

Costs and Benefits of Food Foraging for a Braconid Parasitoid

The costs and benefits of food foraging for the insect parasitoid, Phanerotoma franklini Gahan (Hymenoptera: Braconidae), were examined. Feeding benefited P. franklini by increasing longevity, but did not influence egg maturation. Costs of feeding are represented by the time required to locate and consume food. Field observations of starved wasps revealed that wasps spent approximately 25% of their time grazing on substances on the surface of cranberry foliage. However, our initial laboratory tests to determine the value of these substances were inconclusive. We also examined how grazing behavior integrated with host foraging by comparing the attack rate of fed and unfed wasps held under different host densities. While feeding status did not affect the mean attack rate of wasps, starvation enhanced the probability that a wasp will engage in movement over the host plant foliage. We conclude that feeding benefits P. franklini by increasing longevity, that travel time to food sources is likely to be low, and that the observed food-foraging behavior does not appear to influence host-foraging efficiency.     

Thynnine wasps discriminate among heights when seeking mates: tests with a sexually deceptive orchid

The flower of a sexually deceptive orchid, Chiloglottis reflexa, mimics both the sex pheromone and the appearance of a female thynnine wasp (Neozeloboria nr. proxima). The flower is pollinated when visited by male wasps, who attempt mating with the flower. We have used these mimetic flowers to investigate mating behavior of the male wasps. In field choice experiments, males strongly prefer to visit flowers that are very low in the habitat, 15 cm, vs. flowers that are placed at 55 or 105 cm. These studies suggest that male precopulatory response is strongly dependent on the microlocation of the female (or female mimic). Other insect-mimicking orchids, which together attract several groups of Hymenoptera, may be useful in analogous experiments on mating behavior. Additionally, these experiments help elucidate features of the mimetic flowers, particularly stature, that act to efficiently attract potential pollinators.     

Uscana lariophaga, West-African egg parasitoid of the cowpea bruchid beetleCallosobruchus maculatus: photoperiod, parasitization and eclosion interactions

The diurnal pattern of parasitization and eclosion of the trichogrammatidUscana lariophaga Steffan, egg parasitoid of the cowpea bruchidCallosobruchus maculatus (Fabricius) was studied for L12:D12 photoperiods. The wasp parasitized eggs throughout each 24-h period with about 70% of the parasitization taking place during the first 12 h regardless whether it was the photophase or the scotophase. More female progeny was produced when the first 12-h period was photophase instead of scotophase. Eclosion of wasps took place over a period of 4 days and occurred during the second half of the scotophase and the first half of the photophase. The number of wasps eclosed during the photophase was similar to that eclosed during the scotophase. Average development time was 8.9 days at 30°C. Male development was completed 6 to 8 h before the females, hence a higher percentage of females emerged in the later eclosion peaks. During scotophases more females eclosed than during photophases. Although the results indicate that the wasp is able to perform under dark storage conditions the effect of permanent low light intensity remains to be studied.     

Weak interactions between avian and insect frugivores: the case of Pistacia terebinthus L. (Anacardiaceae)

Vertebrate frugivores often feed on fruits upon or within which insects also feed, yet little information exists on the potential magnitude of interactions between these consumers. The Mediterranean shrub Pistacia terebinthus, the birds that consume its fruits, and the wasps that feed upon its seeds are examined in this study. P. terebinthus produces a highly variable fraction of final-sized red fruits that never become mature (green-colored). Red fruits can be immature, parthenocarpic, aborted, or attacked by wasps, and their pulp is much less nutritious than that of mature fruits. A total of 20 bird species consumed the fruits in the study area. Legitimate dispersers accounted for 39% of the total fruit removal, while pulp eaters and seed predators accounted for the remainder. Birds strongly preferred the mature fruits (only 4% of the fruits consumed were red). The incidence of wasps in the seeds ranged from 0 to 42% of the crop in 1989 and from 0 to 24% in 1990. The influence of avian and insect frugivore guilds on each other appears to be quite low because of the narrow overlap in resource utilization by birds and wasps, and an overall low intensity of wasp seed predation. From an evolutionary perspective, the possible ability of wasps to preclude fruit maturation appears not to be attributable to the present interaction with avian frugivores.     

Transmission of Podisus maculiventris tremulatory signals through plants

Males of the predaceous stink bug Podisus maculiventris (Say) (Heteroptera: Pentatomidae: Asopinae) emit low frequency tremulatory signals. Laser vibrometry was used to record and analyze naturally emitted signals, focusing on variation in signal velocity and frequency during transmission through plants (Phaseolus vulgaris L. and Plumbago auriculata Lam.) as a function of distance from the vibrational source. Signal velocity varied individually between 2 and 15 mm/s recorded on a plant close to the calling male and decreased by 0.3 to 1.5 dB/cm on bean and 0.3 to 0.9 dB/cm on plumbago. The dominant frequency of signals was variable at frequencies below 50 Hz. On bean frequencies centered around 10 Hz or 20 Hz were dominant for signals recorded at the source. Transmission through bean resulted in an increase in the 20 Hz peak relative to other frequencies in the signal. Variation of the dominant frequencies of signals transmitted through plumbago stems were more predictable, showing typical changes in amplitude relative to the distance from the source. The regular variation of the dominant frequency along the stem with linear increase of signal velocity at decreasing distance from the source may provide plant-dwelling insects with information about the distance to the calling individual.     

The aerodynamic signature of running spiders

Many predators display two foraging modes, an ambush strategy and a cruising mode. These foraging strategies have been classically studied in energetic, biomechanical and ecological terms, without considering the role of signals produced by predators and perceived by prey. Wolf spiders are a typical example; they hunt in leaf litter either using an ambush strategy or by moving at high speed, taking over unwary prey. Air flow upstream of running spiders is a source of information for escaping prey, such as crickets and cockroaches. However, air displacement by running arthropods has not been previously examined. Here we show, using digital particle image velocimetry, that running spiders are highly conspicuous aerodynamically, due to substantial air displacement detectable up to several centimetres in front of them. This study explains the bimodal distribution of spider's foraging modes in terms of sensory ecology and is consistent with the escape distances and speeds of cricket prey. These findings may be relevant to the large and diverse array of arthropod prey-predator interactions in leaf litter.     

The halteres of the blowfly Calliphora

The movement of the halteres during fixed flight was video recorded under stroboscopic illumination phase coupled to the wing beat. The halteres swing in a rounded triangular manner through an angle of almost 80° in vertical planes tilted backwards from the transverse plane by ca. 30° (Figs. 1, 2).The physics of the halteres are described in terms of a general formula for the force acting onto the endknob of the moving haltere during rotations and linear accelerations of the fly (Eq. 1). On the basis of the experimentally determined kinematics of the haltere, the primary forces and the forces dependent on angular velocity and on angular acceleration are calculated (Figs. 3, 4).Three distinct types of angular velocity dependent (Coriolis) forces are generated by rotations about 3 orthogonal axes. Thus, in principle one haltere could detect all rotations in space (Fig. 6).The angular acceleration dependent forces have the same direction and frequency as the Coriolis forces, but they are shifted in phase by 90°. Thus, they could be evaluated in parallel and independently from the Coriolis forces. They are, however, much smaller than the Coriolis forces for oscillation frequencies of the fly up to 20 Hz (Fig. 5). From these considerations it is concluded that Coriolis forces play the major role in detecting body rotations.     

Dispersal of adult female fig wasps 1. Arrivals and departures

Ficus burtt-davyi, like most other fig species (Ficus, Moraceae), is exclusively pollinated by its own unique species of fig wasp, in this caseElisabethiella baijnathi (Chalcidoidea, Agaonidae). Because fig crop development on any one tree is usually synchronised, the small and short-lived female wasps have to migrate and find other trees bearing figs which are at suitable stage of development for oviposition. However, the likelihood of successful location and subsequent arrival at a new host tree is dependent on distance and the effect of environmental factors such as wind and temperature. This study examines the relationship between ambient temperatures and the timing of fig wasps emergence from their natal figs and the commencement of their dispersal flight. The behaviour of the wasps arriving at figs which were ready to be pollinated was also examined. The female wasps did not appear to distinguish between the figs and other parts of the tree when in flight. However, after landing on the tree their search for figs was more directed as they visited more figs than leaves. Short-range recognition of figs appears to be by contact chemo-reception, but the wasps showed a preference for entering figs which did not already contain a female wasp.     

Fig volatiles: Their role in attracting pollinators and maintaining pollinator specificity

Each fig tree species (Ficus) is totally dependent on a specific species of wasp for pollination and the larvae of these wasps only develop in the ovules of their specificFicus host. Because the fig crop on any particular tree is generally highly synchronized, the shortlived female wasps must leave their natal tree in order to find figs which are suitable for oviposition. Chemical volatiles produced by figs when they are ready for pollination are thought to be the means by which the wasps detect a suitable host. Gas chromatograms of the fig volatiles of 7 species ofFicus showed them to be species specific. Age related changes in the volatile profiles were noted as extra volatiles are produced when the figs were ready for pollination.     

The effect of insecticides on the distribution of foracing parasitoids,Diaeretiella rapae [Hym.: Braconidae] on plants

Foraging aphid parasitoids,Diaeretiella rapae M'Intosh, were exposed to sublethal doses of the insecticides pirimicarb, permethrin and malathion on brusslls sprouts plants. Observations on wasp distribution over time revealed that wasps spent less time on sprayed plants, relative to controls and, while on these plants, tended to concentrate activity on unsprayed surfaces. For permethrin and malathion, pesticide residues reversed the stereotypic upward foraging pattern of the wasp. Negative consequences of sublethal pesticide doses for parasitoid foraging efficiency are discussed.      

Hearing in honeybees: localization of the auditory sense organ

Airborne sound signals emitted by dancing honeybees (Apis mellifera) contain information about the locations of food sources. Honeybees can perceive these near field sounds and rely on them to decode the messages of the dance language. The dance sound is characterized by rhythmical air particle movement of high velocity amplitudes. The aim of the present study was to identify the sensory structures used to detect near field sound signals. In an operant conditioning experiment, bees were trained to respond to sound. Ablation experiments with these trained bees revealed that neither mechanosensory hairs on the antennae or head nor bristle fields at the joints of the antenna, but Johnston's organ, a chordotonal organ in the pedicel of the antenna, is used to detect near field sound in honeybees.