Laudatio für Herrn Dr. Martin Kaltenpoth : anlässlich der Verleihung des Förderpreises der Ingrid Weiss / Horst Wiehe Stiftung durch der Deutschen Gesellschaft für allgemeine und angewandte Entomologie am 27. Februar 2007 in Innsbruck

Jeder der Anwesenden kennt die Faszination, die von den Leistungen und der Vielgestaltigkeit von Insekten ausgeht. Insekten bieten eine Fülle von Beispielen für die Evolution völlig neuartiger Strukturen, Verhaltensweisen, Verteidigungsmechanismen usw. Man denke z.B. an die Evolution der Flügel, die Tanzsprache der Bienen, die Evolution hochkomplexen Sozialverhaltens oder das chemische Fachwissen der Bombardierkäfer. Besonders faszinierend wird es dort, wo Insekten mit anderen Organismen interagieren. Berühmt geworden sind die z. B. die symbiotischen Beziehungen zwischen verschiedenen Taxa von Ameisen und Pflanzen oder zwischen höheren Termiten und ihren Pilzgärten. Während Mikroorganismen meist als Krankheitserreger oder Konkurrenten für Insekten auftreten, gibt es auch einige Fälle in denen sich aus diesen negativen Beziehungen positive, ja sogar obligate mutualistische Beziehungen entwickelt haben. Dazu gehören die Darmsymbionten die von vielen Herbivoren „adoptiert“ wurden, um ihre nährstoffarme pflanzliche Nahrung besser ausnutzen zu können. Wir haben es einem der diesjährigen Preisträger, Herrn Dr. Martin Kaltenpoth, zu verdanken, dass wir seit kurzem ein weiteres aufregendes und völlig überraschendes Beispiel für eine Symbiose zwischen einem Insekt und einem Bakterium kennen

The odor of origin: kinship and geographical distance are reflected in the marking pheromone of male beewolves (Philanthus triangulum F., Hymenoptera, Crabronidae)

<p>Abstract</p> <p>Background</p> <p>Pheromones play an important role for mate finding and courtship in many insects. In species where males are the signaling sex, females are expected to choose among potential mates with regard to the emitter's quality and/or genetic compatibility. One important aspect is the balance between negative and positive effects of in- vs. outbreeding. In the present study, we aimed to assess the potential of the territory marking pheromone of European beewolves as an indicator for genetic compatibility in the context of female choice.</p> <p>Results</p> <p>We analyzed the sex pheromone composition of male European beewolves (<it>Philanthus triangulum </it>F., Hymenoptera, Crabronidae) from eight different locations across Central Europe (six in Germany, one in England, and one in Italy). The pheromone constitutes a complex blend of various long-chain hydrocarbons (alkanes, alkenes, alcohols, ketones, and a carbon acid). We demonstrate that pheromone composition differs significantly among distant populations (regional scale), among subpopulations (local scale) and between families within subpopulations. The differences in the pheromone blend are positively correlated with geographical distances as might be expected according to an isolation-by-distance model. On a local scale, family membership has a larger effect on pheromone composition than subpopulation affiliation, while the reverse is true for the regional scale.</p> <p>Conclusion</p> <p>Our results show that male pheromones can contain information on both kinship and geographical origin that may be used by females to choose adaptively among potential mates on the basis of their genetic distance.</p

Larval Rearing Temperature Influences Amount and Composition of the Marking Pheromone of the Male Beewolf, Philanthus triangulum

Pheromones play an important role for courtship and mating in many insect species, and they are shaped by a complex interaction of genetic and environmental factors. Developmental temperature is known to have a strong influence on adult life history, morphology, and physiology, but little is known about its effect on pheromone characteristics. In the present study, the influence of temperature during larval development on the amount and composition of the complex marking pheromone from the cephalic glands of the adult male beewolf, Philanthus triangulum F. (Hymenoptera: Crabronidae), was investigated. Additionally, the effects of temperature on several life-history traits were examined. European beewolf larvae were reared at three constant temperatures (20, 25, and 30° C). Males reared at 20° C showed longer development times and higher mortality, suggesting that low temperatures constitute stressful conditions for developing larvae. After eclosion, the amount and composition of the scent marking secretion of the adult males was analyzed by coupled gas chromatography-mass spectrometry. Males that had been reared at 20° C had significantly less secretion than individuals reared under warmer conditions (25° C and 30° C). Furthermore, larval rearing temperature had a significant effect on the composition of the adult males' pheromone gland content, with warmer rearing conditions leading to higher relative amounts of compounds with high molecular weight. The results show that the temperature during larval development significantly affected the amount and composition of the content of the male pheromone glands, probably due to physiological constraints and competing processes for limited energetic resources. Thus, the pheromone gland content may contain information on developmental conditions of males, which may have consequences for female mate choice decisions and male reproductive success

Adaptive dynamic resource allocation in annual eusocial insects: Environmental variation will not necessarily promote graded control

Background: According to the classical model of Macevicz and Oster, annual eusocial insects should show a clear dichotomous "bang-bang" strategy of resource allocation; colony fitness is maximised when a period of pure colony growth (exclusive production of workers) is followed by a single reproductive period characterised by the exclusive production of sexuals. However, in several species graded investment strategies with a simultaneous production of workers and sexuals have been observed. Such deviations from the "bang-bang" strategy are usually interpreted as an adaptive (bet-hedging) response to environmental fluctuations such as variation in season length or food availability. To generate predictions about the optimal investment pattern of insect colonies in fluctuating environments, we slightly modified Macevicz and Oster's classical model of annual colony dynamics and used a dynamic programming approach nested into a recurrence procedure for the solution of the stochastic optimal control problem. Results: 1) The optimal switching time between pure colony growth and the exclusive production of sexuals decreases with increasing environmental variance. 2) Yet, for reasonable levels of environmental fluctuations no deviation from the typical bang-bang strategy is predicted. 3) Model calculations for the halictid bee Lasioglossum malachurum reveal that bet-hedging is not likely to be the reason for the graded allocation into sexuals versus workers observed in this species. 4) When environmental variance reaches a critical level our model predicts an abrupt change from dichotomous behaviour to graded allocation strategies, but the transition between colony growth and production of sexuals is not necessarily monotonic. Both, the critical level of environmental variance as well as the characteristic pattern of resource allocation strongly depend on the type of function used to describe environmental fluctuations. Conclusion: Up to now bet-hedging as an evolutionary response to variation in season length has been the main argument to explain field observations of graded resource allocation in annual eusocial insect species. However, our model shows that the effect of moderate fluctuations of environmental conditions does not select for deviation from the classical bang-bang strategy and that the evolution of graded allocation strategies can be triggered only by extreme fluctuations. Detailed quantitative observations on resource allocation in eusocial insects are needed to analyse the relevance of alternative explanations, e.g. logistic colony growth or reproductive conflict between queen and workers, for the evolution of graded allocation strategies

The evolution of activity breaks in the nest cycle of annual eusocial bees: a model of delayed exponential growth

BACKGROUND: Social insects show considerable variability not only in social organisation but also in the temporal pattern of nest cycles. In annual eusocial sweat bees, nest cycles typically consist of a sequence of distinct phases of activity (queen or workers collect food, construct, and provision brood cells) and inactivity (nest is closed). Since the flight season is limited to the time of the year with sufficiently high temperatures and resource availability, every break reduces the potential for foraging and, thus, the productivity of a colony. This apparent waste of time has not gained much attention. RESULTS: We present a model that explains the evolution of activity breaks by assuming differential mortality during active and inactive phases and a limited rate of development of larvae, both reasonable assumptions. The model predicts a systematic temporal structure of breaks at certain times in the season which increase the fitness of a colony. The predicted pattern of these breaks is in excellent accordance with field data on the nest cycle of the halictid Lasioglossum malachurum. CONCLUSION: Activity breaks are a counter-intuitive outcome of varying mortality rates that maximise the reproductive output of primitively eusocial nests

Flagellar sensilla in male and female European beewolves, Philanthus triangulum F. (Hymenoptera: Sphecidae)

We investigated the morphology of the antennal sensilla of a sphecid wasp, the European beewolf Philanthus triangulum, to provide an inventory for the species and to compare the sensillar equipment between the sexes. The density of sensilla increased from the base to the tip of the antennae. We distinguished nine different types of sensilla. One type has not yet been described in Hymenoptera. The large sensilla basiconica occurred only on the antennae of female beewolves. We discuss the functional significance of the difference between the sexes and compare our results with data from other sphecids and the honeybee Apis mellifera

Sexual selection and the evolution of male pheromone glands in philanthine wasps (Hymenoptera, Crabronidae)

Background: Sexual selection is thought to promote evolutionary changes and diversification. However, the impact of sexual selection in relation to other selective forces is difficult to evaluate. Male digger wasps of the tribe Philanthini (Hymenoptera, Philanthinae) scent mark territories to attract receptive females. Consequently, the organs for production and storage of the marking secretion, the mandibular gland (MG) and the postpharyngeal gland (PPG), are subject to sexual selection. In female Philanthini, these glands are most likely solely subject to natural selection and show very little morphological diversity. According to the hypothesis that sexual selection drives interspecific diversity, we predicted that the MG and PPG show higher interspecific variation in males than in females. Using histological methods, 3D-reconstructions, and multivariate statistical analysis of morphological characters, we conducted a comparative analysis of the MG and the PPG in males of 30 species of Philanthini and three species of the Cercerini and Aphilanthopsini, two related tribes within the Philanthinae. Results: We found substantial interspecific diversity in gland morphology with regard to gland incidence, size, shape and the type of associated secretory cells. Overall there was a phylogenetic trend: Ensuing from the large MGs and small PPGs of male Cercerini and Aphilanthopsini, the size and complexity of the MG was reduced in male Philanthini, while their PPG became considerably enlarged, substantially more complex, and associated with an apparently novel type of secretory cells. In some clades of the Philanthini the MG was even lost and entirely replaced by the PPG. However, several species showed reversals of and exceptions from this trend. Head gland morphology was significantly more diverse among male than among female Philanthinae. Conclusion: Our results show considerable variation in male head glands including the loss of an entire gland system and the evolution of a novel kind of secretory cells, confirming the prediction that interspecific diversity in head gland morphology is higher in male than in female Philanthini. We discuss possible causes for the remarkable evolutionary changes in males and we conclude that this high diversity has been caused by sexual selection

A cuckoo in wolves' clothing? Chemical mimicry in a specialized cuckoo wasp of the European beewolf (Hymenoptera, Chrysididae and Crabronidae)

<p>Abstract</p> <p>Background</p> <p>Host-parasite interactions are among the most important biotic relationships. Host species should evolve mechanisms to detect their enemies and employ appropriate counterstrategies. Parasites, in turn, should evolve mechanisms to evade detection and thus maximize their success. Females of the European beewolf (<it>Philanthus triangulum</it>, Hymenoptera, Crabronidae) hunt exclusively honeybee workers as food for their progeny. The brood cells containing the paralyzed bees are severely threatened by a highly specialized cuckoo wasp (<it>Hedychrum rutilans</it>, Hymenoptera, Chrysididae). Female cuckoo wasps enter beewolf nests to oviposit on paralyzed bees that are temporarily couched in the nest burrow. The cuckoo wasp larva kills the beewolf larva and feeds on it and the bees. Here, we investigated whether <it>H. rutilans </it>evades detection by its host. Since chemical senses are most important in the dark nest, we hypothesized that the cuckoo wasp might employ chemical camouflage.</p> <p>Results</p> <p>Field observations suggest that cuckoo wasps are attacked by beewolves in front of their nest, most probably after being recognized visually. In contrast, beewolves seem not to detect signs of the presence of these parasitoids neither when these had visited the nest nor when directly encountered in the dark nest burrow.</p> <p>In a recognition bioassay in observation cages, beewolf females responded significantly less frequently to filter paper discs treated with a cuticular extract from <it>H. rutilans </it>females, than to filter paper discs treated with an extract from another cuckoo wasp species (<it>Chrysis viridula</it>). The behavior to paper discs treated with a cuticular extract from <it>H. rutilans </it>females did not differ significantly from the behavior towards filter paper discs treated with the solvent only.</p> <p>We hypothesized that cuckoo wasps either mimic the chemistry of their beewolf host or their host's prey. We tested this hypothesis using GC-MS analyses of the cuticles of male and female beewolves, cuckoo wasps, and honeybee workers. Cuticle extracts of <it>Hedychrum nobile </it>(Hymenoptera: Chrysididae) and <it>Cerceris arenaria </it>(Hymenoptera: Crabronidae) were used as outgroups. There was little congruence with regard to cuticular compounds between <it>H. rutilans </it>females and honeybees as well as females of <it>C. arenaria </it>and <it>H. nobile</it>. However, there was a considerable similarity between beewolf females and <it>H. rutilans </it>females. Beewolf females show a striking dimorphism regarding their cuticular hydrocarbons with one morph having (<it>Z</it>)-9-C25:1 and the other morph having (<it>Z</it>)-9-C27:1 as the major component. <it>H. rutilans</it> females were more similar to the morph having (Z)-9-C27:1 as the main component.</p> <p>Conclusion</p> <p>We conclude that <it>H. rutilans </it>females closely mimic the composition of cuticular compounds of their host species <it>P. triangulum</it>. The occurrence of isomeric forms of certain compounds on the cuticles of the cuckoo wasps but their absence on beewolf females suggests that cuckoo wasps synthesize the cuticular compounds rather than sequester them from their host. Thus, the behavioral data and the chemical analysis provide evidence that a specialized cuckoo wasp exhibits chemical mimicry of the odor of its host. This probably allows the cuckoo wasp to enter the nest with a reduced risk of being detected by olfaction and without leaving traitorous chemical traces.</p

Nitric oxide radicals are emitted by wasp eggs to kill mold fungi

Detrimental microbes caused the evolution of a great diversity of antimicrobial defenses in plants and animals. Insects developing underground seem particularly threatened. Here we show that the eggs of a solitary digger wasp, the European beewolf Philanthus triangulum, emit large amounts of gaseous nitric oxide (NO center dot) to protect themselves and their provisions, paralyzed honeybees, against mold fungi. We provide evidence that a NO-synthase (NOS) is involved in the generation of the extraordinary concentrations of nitrogen radicals in brood cells (similar to 1500 ppm NO center dot and its oxidation product NO2 center dot). Sequencing of the beewolf NOS gene revealed no conspicuous differences to related species. However, due to alternative splicing, the NOS-mRNA in beewolf eggs lacks an exon near the regulatory domain. This preventive external application of high doses of NO center dot by wasp eggs represents an evolutionary key innovation that adds a remarkable novel facet to the array of functions of the important biological effector NO center dot

Host Plant Use by Competing Acacia-Ants: Mutualists Monopolize While Parasites Share Hosts

Protective ant-plant mutualisms that are exploited by non-defending parasitic ants represent prominent model systems for ecology and evolutionary biology. The mutualist Pseudomyrmex ferrugineus is an obligate plant-ant and fully depends on acacias for nesting space and food. The parasite Pseudomyrmex gracilis facultatively nests on acacias and uses host-derived food rewards but also external food sources. Integrative analyses of genetic microsatellite data, cuticular hydrocarbons and behavioral assays showed that an individual acacia might be inhabited by the workers of several P. gracilis queens, whereas one P. ferrugineus colony monopolizes one or more host trees. Despite these differences in social organization, neither of the species exhibited aggressive behavior among conspecific workers sharing a tree regardless of their relatedness. This lack of aggression corresponds to the high similarity of cuticular hydrocarbon profiles among ants living on the same tree. Host sharing by unrelated colonies, or the presence of several queens in a single colony are discussed as strategies by which parasite colonies could achieve the observed social organization. We argue that in ecological terms, the non-aggressive behavior of non-sibling P. gracilis workers — regardless of the route to achieve this social structure — enables this species to efficiently occupy and exploit a host plant. By contrast, single large and long-lived colonies of the mutualist P. ferrugineus monopolize individual host plants and defend them aggressively against invaders from other trees. Our findings highlight the necessity for using several methods in combination to fully understand how differing life history strategies affect social organization in ants