Influence of temperature on host location and multisensory orientation in the parasitoid Pimpla turionellae (L.) (Hymenoptera)

The ichneumonid wasp Pimpla turionellae (L.) (Hymenoptera) is a specialist parasitoid of lepidopteran pupae, and has to overcome the challenge of reduced chemical and visual cues as pupae are immobile, do not feed and do not emit excrements. Certain hymenopteran species have developed a particular mechanosensory mechanism in order to locate hosts hidden in hollow spaces inside of plant material (Broad & Quicke 2000). Similar to echolocation they use self-produced vibrations, instead of sounds, that are transmitted by the antennae onto the substrate. In analogous way to acoustics, this mechanosensory mechanism is referred to as vibrational sounding (Wäckers & al. 1998). Thermal dependence is well known in acoustical and vibrational communication of arthropods (e.g. Pires & Hoy 1992, Shimizu & Barth 1996) and is likewise presumed to affect mechanosensory host location by vibrational sounding. The species P. turionellae has recently been found to use vibrational sounding successfully in a temperature range from 8 to 28°C, but with less performance of searching behaviour and an adjusted signal production at extreme temperatures (Kroder & al. 2006, & al. 2007b, Samietz & al. 2006). Many insects have evolved strategies to maintain a balance of body temperature by ecto- and endothermic means. Raising and maintaining body temperatures above the ambient environment by endothermic means is particularly known in several hymenopteran species (Heinrich 1993). In the case of a thermally influenced host location mechanism, such means of thermoregulation could be supposed as well in order to maintain performance with changing temperatures. The study elucidates if the wasps are able to regulate their body temperature at suboptimal conditions during vibrational sounding and furthermore examines the role of vibrational sounding in multisensory orientation at different ambient temperatures.Die Schlupfwespe Pimpla turionellae parasitiert versteckte Lepidopterenpuppen und orientiert sie sich bei der Wirtssuche multisensorisch mittels visueller Reize und aktiver Vibrationsortung mit selbst produzierten Schwingungen (Vibrational-Sounding). Die Studie untersucht, inwieweit die Wespen bei Änderung der Umgebungstemperaturen von 8-26°C (1) zwischen der temperatur-sensitiven vibratorischen und der -insensitiven visuellen Orientierung wechseln und (2) gegebenenfalls selbst die Körpertemperatur regulieren können, um die sehr präzise Vibrationsortung bei niedrigen Temperaturen aufrechtzuerhalten. Messungen mit Infrarot-Thermographie zeigen, dass suchende Wespen leicht erhöhte Körpertemperaturen während der vibratorischen Wirtssuche aufweisen, welche auf metabolische Wärmeproduktion zurückzuführen sind. Wahlexperimente unter kontrollierten Temperaturen zeigen zudem, dass die Nutzung der temperatur-sensitiven vibratorischen Reize bei pessimalen Temperaturen abnimmt und die Wespen auf fast ausschließliche visuelle Orientierung wechseln. Folglich wird die Relevanz einzelner Reize bei der multisensorischen Orientierung direkt vom Faktor Temperatur beeinflusst. Solange ein zuverlässiger Reiz vorhanden ist, nimmt dabei auch die Präzision der Lokalisation insgesamt nicht ab

A female-specific attractant for the codling moth, Cydia pomonella , from apple fruit volatiles

Host plant-derived esters were investigated as potential female-specific attractants for the codling moth (CM), Cydia pomonella (L.), a key pest of apples worldwide. The behavioural effects of single and combined volatile compounds and of a natural odour blend were examined using olfactometry and wind-tunnel bioassays. The apple-derived volatile butyl hexanoate attracted mated females while it was behaviourally ineffective for males over a dosage range of more than three orders of magnitude in olfactometer assays. Female CM preferred this kairomone to the headspace volatiles from ripe apples. Both no-choice and choice trials in the wind-tunnel suggested that female moths might be effectively trapped by means of this compound. In contrast, headspace volatiles collected from ripe apple fruits as well as a blend containing the six dominant esters from ripe apples were behaviourally ineffective. A female-specific repellency was found for the component hexyl acetate in the olfactometer, but this ester had no significant effect in the wind-tunnel. Butyl hexanoate with its sex-specific attraction should be further evaluated for monitoring and controlling CM females in orchard

Relationship Between Behavior and Physiology in an Invasive Pest Species: Oviposition Site Selection and Temperature-Dependent Development of the Oriental Fruit Moth (Lepidoptera: Tortricidae)

Oviposition site selection is crucial for the reproductive success of a herbivore insect species with relatively sedentary larvae. The optimal oviposition theory, i.e., the preference-performance hypothesis, has thus far mainly been tested with a focus on nutritional quality of the host. This study investigates whether female oriental fruit moth Grapholita (Cydia) molesta choose a microhabitat for oviposition characterized by a temperature range within which their offspring perform best. Thermal preferences of females during oviposition were assessed in a circular temperature gradient arena. Offspring performance and survival were assessed under different constant temperature conditions. Females preferred oviposition sites of ∼30°C over lower and higher temperatures. At this temperature, egg, larval, and pupal development was significantly faster than at 22 and 25°C, and larval development was also faster than at 33°C. At 30°C and at the lower temperatures tested, survival of eggs and larvae was significantly higher than at 33°C, whereas development was precluded at 35°C. Furthermore, female pupal weight attained at 30 and 33°C exceeded that reached at the lower temperatures tested. Considering the potentially reduced predation risk caused by the shorter developmental time of eggs and larvae, the laboratory data suggest that this species maximizes its fitness by selecting a thermally optimal environment for its offspring, supporting the optimal oviposition theory. Conversely, it is known that the codling moth (C. pomonella) lacks a mechanism to avoid temperatures lethal to progeny development, which may reflect the differences in geographic ranges of these tortricid

Host recognition in a pollen-specialist bee: evidence for a genetic basis

To investigate the effect of larval pollen diet on floral choice in a specialized bee species, we compared the floral preferences of individuals of Heriades truncorum (Megachilidae) reared on host pollen with those of individuals reared on two different types of non-host pollen. Females were allowed to nest in cages where both host and non-host flowers were available. All females, regardless of larval diet, restricted pollen collection to their host, although they visited the flowers of both host and non-host plants for nectar. When offered only the non-host pollen source, females ceased nesting activities. Males reared on non-host pollen exclusively restricted their patrolling flights to flowers of their normal host. This study provides the first empirical investigation of the imprinting theory in oligolectic bees, and unambiguously suggests that host recognition has a genetic basis in H. truncorum. We discuss the implication of this finding for the understanding of bee-flower relationship

Temperature affects interaction of visual and vibrational cues in parasitoid host location

Parasitoid host location in nature is facilitated by simultaneously using different information sources. How multisensory orientation on the same spatial scale is influenced by environmental conditions is however poorly understood. Here we test whether changes in reliability of cues can cause parasitoids to alter multisensory orientation and to switch to cues that are more reliable under extreme temperatures. In the ichneumonid wasp Pimpla turionellae, multisensory use of thermally insensitive vision and thermally sensitive mechanosensory host location by vibrational sounding (echolocation on solid substrate) was investigated with choice experiments on plant-stem models under optimum temperature (18°C), at high- (28°C) and low-temperature limits (8°C) of vibrational sounding. Temperature affected relative importance of vibrational sounding whereas visual orientation did not vary. At 18°C, parasitoids used visual and vibrational cues with comparable relative importance. At 8 and 28°C, the role of vibrational sounding in multisensory orientation was significantly reduced in line with decreased reliability. Wasps nearly exclusively chose visual cues at 8°C. The parasitoids switch between cues and sensory systems depending on temperature. As overall precision of ovipositor insertions was not affected by temperature, the parasitoids fully compensate the loss of one cue provided another reliable cue is available on the same spatial scal

No kin discrimination in female mate choice of a parasitoid with complementary sex determination

Discrimination against kin as mates, via genetic or environmentally derived cues of relatedness, can prevent inbreeding and thus enhance individual fitness and promote population survival. Sex in the parasitoid wasp Cotesia glomerata L. (Hymenoptera: Braconidae) is determined by one locus with multiple alleles, a mechanism termed single-locus complementary sex determination (sl-CSD). Under sl-CSD, haploid individuals are males, whereas diploid individuals are females when heterozygous at the sex determination locus but males when homozygous. In species with sl-CSD, inbreeding leads to increased incidence of matings between individuals sharing an allele at the sex locus and thus to increased diploid male production. Diploid males cause an undesirable sex ratio distortion and can be of inferior fitness. To evade these deleterious effects, species with sl-CSD are expected to avoid inbreeding. We investigated whether C. glomerata females discriminate against close kin as mating partners. We performed a mate choice experiment, which allowed us to distinguish between kin discrimination based on the perception of phenotype-related cues and kin discrimination based on the perception of cues associated with the developmental environment. As kin discrimination is often mediated through cuticular hydrocarbons (CHCs), we additionally examined composition of the CHC profiles of males. We found no evidence for discrimination against related or familiar males nor for differences in the CHC profiles of males. These results indicate that kin discrimination is not a relevant inbreeding avoidance strategy in C. glomerat

Host Location of a Pupal Parasitoid in a Tritrophic System Compared to a Model Offering Mechanosensory Cues Only

Several species of hymenopteran parasitoids are able to locate concealed pupal hosts by vibrational sounding. However, the specific role of this technique of mechanosensory host location has not yet been investigated in a natural, tritrophic system with multiple cues. Here we compared the host location of the pupal parasitoid Xanthopimpla stemmator in a tritrophic system with corn borer pupae in maize stem to the behavior on a paper model offering mechanosensory cues only. In general, the behavioral pattern and the behavioral states exhibited by host-searching female parasitoid were identical in the model and in the tritrophic system, while quantitative aspects differed. Our results demonstrate that vibrational sounding maintains its significance for host location in an environment with multiple cues, and that additional cues may increase responsiveness of female

Consequences of mixed species infestation on the searching behavior and parasitism success of a larval parasitoid

When two herbivore pest species are potential hosts of a single parasitoid species, two questions arise. Firstly, which host is preferable for mass rearing in terms of later parasitoid performance, and secondly, how do parasitoids perform in mixed herbivore situations after colony establishment? We tested Hyssopus pallidus, a gregarious parasitoid of two major pests of apple, Cydia (Grapholita) molesta and Cydia pomonella, before and after landing on apples infested by one of the two Cydia species. Pre-alighting host preference was tested in a Y-tube olfactometer setup, and parasitism success in a contact bioassay. To gain information on parasitoid performance throughout the growing season, different fruit growth stages were used. Irrespective of the host they had developed on, the parasitoids showed similar olfactory preferences when given a dual choice between infested and healthy fruits, and they did not discriminate between fruits infested by C.molesta and C.pomonella. Responsiveness was generally high, especially late in the season close to harvest. Both hosts are parasitized regardless of the host the parasitoid female had developed on, and no differences in parasitism rates or number of offspring were noted for the two hosts offered. Results were consistent for all apple growth stages tested. In conclusion, mass rearing of this parasitoid can be carried out on either host, without limiting the future efficacy of the bio-control agent. Similarly, established colonies are expected to develop further on both hosts without any bias in host preferenc

Visual Orientation of Overwintered Anthonomus pomorum (Coleoptera: Curculionidae)

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 fiel

Ability of the Oriental Fruit Moth Grapholita molesta (Lepidoptera: Tortricidae) to Detoxify Juglone, the Main Secondary Metabolite of the Non-host Plant Walnut

Many plant species produce toxic secondary metabolites that limit attacks by herbivorous insects, and may thereby constrain insect expansion to new hosts. Walnut is a host for the codling moth Cydia pomonella, which efficiently detoxifies the main walnut defensive compound juglone (5-hydroxy-1,4-naphthoquinone). The oriental fruit moth Grapholita molesta, which also belongs to the tribe Grapholitini, does not feed on walnut. We tested the performance of G. molesta, a highly invasive species, on artificial diets containing juglone at levels mimicking those found in walnut over the growing season. Juglone-fed G. molesta survived relatively well to adulthood, but larval and adult body weights were reduced, and larval developmental time was prolonged in a dose-dependent fashion. Chemical analysis of frass from larvae that had been fed a juglone-containing diet suggests that G. molesta reduces juglone to non-toxic 1,4,5-trihydroxynaphthalene in its gut. This unexpected tolerance of G. molesta to high levels of juglone may facilitate expansion of the host range beyond the current rosacean fruit trees used by this invasive pes