Jörg Samietz

Interaction of vibrational and visual cues in parasitoid host location

Abstract. Female parasitoids are guided by multisensory information during host finding. Individual cues are used in an interactive or a hierarchical manner according to the relative importance on the spatial scale of their effect. Unlike most studies that concentrate on single cues, the present paper investigates the interaction of two physical cues. The interaction of mechanosensory and visual cues was studied in the pupal parasitoid Pimpla turionellae (Hymenoptera: Ichneumonidae). This species uses, amongst other senses, vibrational sounding (echolocation in a solid substrate) to find its mainly endophytic hosts. Location and frequency of ovipositor insertions were scored on cylindrical plant stem models with single or combined cues. Single-cue experiments show that parasitoids use both visual and mechanosensory cues and achieve a similar precision of host location with either cue. The combination of vision and vibrational sounding increased the precision of host location by a factor of approximately two to three. We conclude that the two senses interact, resulting in an additive accuracy. Neither the visual nor the mechanosensory cue was favored when offered adjacent to each other on the same stem model. On the investigated spatial scale, both physical cues are used and seem to be equally important for host location in this species.

Thermoregulation behaviour in codling moth larvae

Abstract.  The thermoregulation behaviour of the codling moth, Cydia pomonella, is investigated in temperature gradient experiments with larvae feeding within apples, and with mature larvae searching for cocooning sites. Feeding larvae appear to prefer the apple hemisphere with a higher temperature (i.e. they build larger cavities in the radiated, warmer part of the fruit). The proportion of larval cavities in the warmer hemisphere is positively related to increasing apple temperature on that side, as well as to the temperature difference between the warm and the cool fruit hemisphere. The mechanism in feeding larvae can be termed as cryptic basking because, during microhabitat selection, the caterpillars exploit temperature differences that are caused explicitly by incident solar radiation. Fifth‐instar larvae in search of cocooning sites show no temperature preference within the large gradient offered (9–29 °C), with no difference between males and females. During larval development, the insect changes its thermoregulation behaviour in response to a possible shift in benefits of an elevated body temperature with respect to environmental conditions. Both the thermoregulation behaviour and such a shift of behavioural response should be respected when simulating body temperatures of the species.

Visual orientation of overwintered Anthonomus pomorum (Coleoptera: Curculionidae)

Abstract Vision is considered to be an important component of the sensory system used by herbivorous insects to locate host plants. We investigated preferences for transmissive colors and contrasts in the apple blossom weevil, Anthonomus pomorum L., which colonizes apple trees, Malus domestica Borkh., in early spring. The main components of the supposed visual system, green, blue, and UV light, were offered to the weevils versus black or versus each other in a dual choice arena. Furthermore, silhouettes of two different achromatic contrasts were offered in front of the colors blue and green. We also tested whether visual preferences change with time, i.e., 2, 4, and 6 d after termination of diapause in early spring. Female weevils chose UV, green, and blue over black, whereas males only chose UV over black in a series of dual choice experiments. Both sexes preferred blue to green. Weevil response to silhouettes of different contrasts was influenced by the background color, i.e., the silhouette with the higher contrast was preferred in front of green, whereas the silhouette with the lower contrast was preferred in front of blue. The preferences during visual orientation did not change with time after diapause. The more discriminating response of females may reflect a greater visual acuity or a greater visual responsiveness in females than in males because of a higher affinity for the host plant. Our findings indicate that weevils might have a trichromatic visual system and that they are able to include visual cues for the location of host plants in the field.

Perception of Chromatic Cues During Host Location by the Pupal Parasitoid Pimpla turionellae (L.) (Hymenoptera: Ichneumonidae)

Abstract Chromatic and achromatic plant cues are expected to be particularly important for parasitoids of endophytic pupal hosts, because these stages do not feed and therefore avoid volatile emission caused by plant tissue damage. Endophytic feeding can cause discoloration or desiccation, leading to changes in color and/or brightness of infested plant parts that may be visually detected by parasitoids. The role of color cues in the host-finding behavior of parasitoids is poorly understood, and the visual system of most parasitoid species has not yet been investigated. We studied color discrimination ability and innate color preferences in the pupal parasitoid Pimpla turionellae (L.) (Hymenoptera: Ichneumonidae) during location of concealed hosts. Responses to combinations of yellow and blue bands of different reflectance intensities were investigated on cylindrical models of plant stems. The parasitoid’s reaction to these chromatic cues was evaluated by scoring the number of ovipositor insertions into the colored bands. Female parasitoids discriminated blue from yellow irrespective of total reflectance and inserted their ovipositors significantly more often into the blue area. True color vision is demonstrated for the examined species, and responses to chromatic cues are discussed in relation to their importance for host location in parasitoids. Results of this study and of our previous work suggest that P. turionellae uses contrasts (chromatic or achromatic) rather than specific color characteristics in visual host location.

Efficiency of vibrational sounding in parasitoid host location depends on substrate density

Parasitoids of concealed hosts have to drill through a substrate with their ovipositor for successful parasitization. Hymenopteran species in this drill-and-sting guild locate immobile pupal hosts by vibrational sounding, i.e., echolocation on solid substrate. Although this host location strategy is assumed to be common among the Orussidae and Ichneumonidae there is no information yet whether it is adapted to characteristics of the host microhabitat. This study examined the effect of substrate density on responsiveness and host location efficiency in two pupal parasitoids, Pimpla turionellae and Xanthopimpla stemmator (Hymenoptera: Ichneumonidae), with different host-niche specialization and corresponding ovipositor morphology. Location and frequency of ovipositor insertions were scored on cylindrical plant stem models of various densities. Substrate density had a significant negative effect on responsiveness, number of ovipositor insertions, and host location precision in both species. The more niche-specific species X. stemmator showed a higher host location precision and insertion activity. We could show that vibrational sounding is obviously adapted to the host microhabitat of the parasitoid species using this host location strategy. We suggest the attenuation of pulses during vibrational sounding as the energetically costly limiting factor for this adaptation.

The role of the plant in attracting parasitoids: response to progressive mechanical wounding

Based on the model system of Brussels sprouts [Brassica oleracea var. gemmifera (Brassicaceae)], the herbivore cabbage white caterpillar, Pieris brassicae (L.) (Lepidoptera: Pieridae), and the parasitoid wasp, Cotesia glomerata (L.) (Hymenoptera: Braconidae), the influence of plant damage type, and damage duration were assessed on plant volatile emission and subsequent recruitment of natural antagonists of the herbivore. Plants were damaged by three methods for a period of either 3 or 8 h: herbivore damage (HD), progressive mechanical damage, and final mechanical damage inflicted in a single event. Wind‐tunnel bioassays evaluated whether the mode of damage affected female parasitoid oriented flight. After both periods of damage, all treatments were highly significantly preferred by naïve C. glomerata to undamaged control plants. After 3 h, herbivore‐damaged plants were significantly preferred to plants with final damage (FD). Most remarkably, following 8‐h damage, the parasitoid preferred both herbivore‐damaged and progressively damaged plants to plants with FD and did not significantly discriminate between herbivore and progressively damaged plants, thus indicating a similarity in plant response to herbivore and progressive mechanical damage. In addition to wind‐tunnel bioassays, emitted plant volatiles were collected and analysed by thermal desorption gas chromatography/mass spectrometry, following 3 and 8 h of damage in order to correlate volatiles released from different damage types with the attraction of the parasitoid. Differences in volatile profiles from all damage types were similar following both 3 and 8 h of damage, with only (Z)‐3‐hexenyl acetate found to be emitted in significantly higher quantities by final mechanical damage compared with HD after 3 h. In conclusion, the plants response to progressive mechanical damage was more similar to HD than final mechanical damage deployed at a single point in time, irrespective of damage duration, and C. glomerata did not significantly discriminate between progressive damage and HD.

Thermal response in adult codling moth

Abstract.  The thermoregulation behaviour of the adult codling moth, Cydia pomonella, is investigated in the laboratory using temperature gradient experiments. Unmated males and females are tested at dawn when moths typically move to resting sites. Mated females are tested during oviposition over a complete diurnal cycle. Temperature strongly affects microhabitat selection in adult moths. Unmated males and females prefer to rest at the low‐temperature ends of temperature gradients between 15 and 32 °C. Relative humidity does not influence the thermal response in unmated females, whereas males show a less distinct temperature selection under high humidity. By contrast to unmated moths, ovipositing females prove to be highly thermophilous (i.e. they deposit the highest proportions of their eggs in the zones of highest temperatures of gradients between 15 and 36 °C). This striking discrepancy in thermal response of females between their premating and oviposition period is likely to reflect an adaptation to different selection pressures from the thermal environment. Unmated moths may benefit from low temperatures by a longer lifespan and crypsis within the tree canopy, whereas the choice of warmer oviposition sites by mated females will favour a faster development of eggs.

Effect of plant architecture and hail nets on temperature of codling moth habitats in apple orchards

Plant architecture of apple trees in commercial orchards was rapidly changed from traditional tall trees to dwarf trees to optimize yield and fruit quality. Additionally, hail nets are widely used to prevent yield loss by hail. These changes are expected to considerably influence the orchard microclimate and thus the developmental rates of pest insects in apple. However, these relationships have not yet been fully elucidated. The present study was conducted over the seasonal cycle to investigate the influence of plant architecture and hail nets on the habitat temperatures of the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), in apple, Malus domestica Borkh. (Rosaceae). Within the canopies, leaf area index (LAI) and global site factor (GSF) were quantified using hemispherical photography. Temperature was analysed for the main habitats of the different codling moth stages, i.e., air within the canopy, bark of tree stems, and apple fruit. In dwarf trees, LAI was lower, leading to a higher GSF than in tall trees. Hail nets did not influence LAI and GSF. Results for dwarf trees compare as follows with those for tall trees: Average air temperatures within the canopy were 0.7 °C higher during daytime, whereas 0.4 °C lower at night. Mean surface temperatures of bark were 0.9 °C higher on sunny and 0.4 °C on overcast days. Mean surface temperatures of apple fruits were 1.8–2.7 °C higher on sunny days, but 0.6 °C cooler on overcast days. The effect of hail nets was confined to a reduction of the air temperature within the canopy by approximately 0.2–0.8 °C. Bark and apple surface temperatures were not significantly affected. Based on the temperature differences in the habitats considered, the calculated development of the codling moth in dwarf trees was on average 3 days faster than in tall trees. The calculations imply a negligible effect of hail nets on codling moth development.

Temperature effects on egg development of the rosy apple aphid and forecasting of egg hatch

The development of Dysaphis plantaginea (Pass.) (Homoptera: Aphididae) winter eggs was studied at six different constant temperatures ranging from 7.5 to 16.5 °C in order to improve the basis for phenological forecasts in early spring. The mortality was generally low at temperatures below 13.5 °C but increased considerably at 16.5 °C. The effect of temperature on development rates could be described with linear regression within the temperature range under study. The lower temperature threshold for development was estimated to be 4.0 °C and the thermal constant 140 day‐degrees. A time‐varying distributed delay approach was used to establish a temperature driven phenology model for winter egg hatch of D. plantaginea considering the intrinsic variability in development time. The model parameters such as temperature‐dependent development times and corresponding variances were quantified based on the experimental data. When compared with independent observations on egg hatch under semifield conditions, the model gave satisfactory validation results. It can be used as forecasting tool for the optimal timing of monitoring and control measures for D. plantaginea in early spring.

Monitoring the dynamics of orchard colonisation by Anthonomus pomorum in spring

Overwintered adult apple blossom weevils, Anthonomus pomorum (L.) (Coleoptera: Curculionidae), colonise apple trees, Malus domestica Borkh. (Rosaceae), in early spring. Information gained from a suitable monitoring technique could serve as a guide to determining the accurate timing for control measures. To assess tree colonisation by flight and crawling, Plexiglas flight traps and commercial screen traps developed for different curculionid species were used. Refuges were mimicked by a newly designed shelter trap based on transparent bubble wrap. The trap catches were contrasted to the limb jarring technique. Microclimate and weather conditions were determined over a trial period of one and a half months. The shelter traps caught a high number of weevils irrespective of weather conditions, and thus performed better than both flight and screen traps. The temperature level within the shelter traps was slightly, but significantly, higher than on the trunk, indicating that weevils exploit microclimatic differences and occupy thermally favourable refuges. The seasonal culmination of colonisation as determined by shelter traps coincided with that as determined by the two methods used to monitor weevil movement towards host trees (flight traps and screen traps). The data indicated that not only flight, but also crawling as quantified by the screen traps contributes substantially to spring colonisation. Captures by all three types of traps over the spring season were significantly positively correlated with each other, but in most respects not with the results of the limb jarring. We conclude that the newly designed shelter trap will be useful for accurately monitoring the seasonal course of colonisation and holds promise for determining the timing of interventions.