Succession of Dung-Inhabiting Beetles and Flies Reflects the Succession of Dung-Emitted Volatile Compounds.

Chemical cues, such as volatile organic compounds (VOCs), are often essential for insects to locate food. Relative to the volume of studies on the role of VOCs in insect-plant relationships, the role of VOCs emitted by dung and carrion in mediating the behavior of insect decomposers is understudied. Such relationships may provide a mechanistic understanding of the temporal axis of community assembly processes in decomposing insect communities. We focused on the temporal succession of volatiles released by cow dung pats and the potential influence on dung-inhabiting insects. Using gas chromatography/mass spectrometry we identified and quantified VOCs released from dung 1-h, and 1, 2 3, 5, and 7 d-old. We then related changes in VOCs to successional patterns of dung-inhabiting beetles and flies. We detected 54 VOCs which could be assigned to two successional groups, with chemical turnover in dung changing around day 2. The early successional group consisted primarily of aliphatic alcohols and phenols, and the late one of aliphatic esters, nitrogen- and sulfur-bearing compounds. Flies were predominately associated with the early successional group, mainly with 1-butanol. Beetles were associated predominately with the late-successional group, mainly with dimethyl trisulfide. This association between insect and chemical successional patterns supports the idea that habitat filtering drives the community assembly of dung-inhabiting insects on an aging resource. Moreover, the affinity of both insect groups to specific VOC groups provides a mechanistic explanation for the predictability of successional patterns found in dung-inhabiting insect communities.

Distinct Roles of Cuticular Aldehydes as Pheromonal Cues in Two Cotesia Parasitoids.

Cuticular compounds (CCs) that cover the surface of insects primarily serve as protection against entomopathogens, harmful substances, and desiccation. However, CCs may also have secondary signaling functions. By studying the role of CCs in intraspecific interactions, we may advance our understanding of the evolution of pheromonal communication in insects. We previously found that the gregarious parasitoid, Cotesia glomerata (L.), uses heptanal as a repellent pheromone to help avoid mate competition among sibling males, whereas another cuticular aldehyde, nonanal, is part of the female-produced attractive sex pheromone. Here, we show that the same aldehydes have different pheromonal functions in a related solitary parasitoid, Cotesia marginiventris (Cresson). Heptanal enhances the attractiveness of the female's sex pheromone, whereas nonanal does not affect a female's attractiveness. Hence, these common aldehydes are differentially used by the two Cotesia species to mediate, synergistically, the attractiveness of the main constituents of their respective sex pheromones. The specificity of the complete sex pheromone blend is apparently regulated by two specific, less volatile compounds, which evoke strong electroantennographic (EAG) responses. This is the first demonstration that volatile CCs have evolved distinct pheromonal functions to aid divergent mating strategies in closely related species. We discuss the possibility that additional compounds are involved in attraction and that, like the aldehydes, they are likely oxidative products of unsaturated cuticular hydrocarbons.

Honeybee Pollinators Use Visual and Floral Scent Cues to Find Apple (Malus domestica) Flowers.

Apple flowers of most varieties require pollinator-mediated cross-pollination. However, little is known about the cues used by pollinators to find the flowers. We used bioassays to investigate the importance of visual and olfactory cues for the attraction of honeybee pollinators to apple flowers. Chemical-analytical and electrophysiological approaches were used to determine floral scents and investigate antennal responses of honeybees to scents from flowering twigs. Bioassays showed that visual and olfactory cues were equally important for the attraction of honeybees to apple flowers. Floral scents were dominated by aromatic components, mainly benzyl alcohol, and the antennae of honeybees responded to a large number of components, among them to benzyl alcohol, linalool, and indole. Our study aims to better understand how this important fruit crop communicates with its main pollinators. This knowledge might be used to improve the attractiveness of apple flowers to pollinators to optimize fruit sets.

The Combined Use of an Attractive and a Repellent Sex Pheromonal Component by a Gregarious Parasitoid.

Gregarious parasitoids usually clump their cocoons together and the adults emerge in a synchronized fashion. This makes it easy for them to find mating partners and most copulations indeed take place at the natal patch. Yet, males should leave such sites when females are no longer receptive. As yet, this decision-making process and the possible involvement of pheromones were poorly understood. Here we report on a remarkable use of attractive and repellent pheromones of the well-studied gregarious parasitoid species Cotesia glomerata (L.) (Hymenoptera: Braconidae). Virgin C. glomerata females were found to release an attractive as well as a repellent compound, which in combination arrest males on the natal patch, but after mating the females stop the production of the attractant and the males are repelled. The repellent compound was identified as heptanal, which was also released by males, probably reducing male-male competition on the natal patch. We also confirmed that the sex ratio of the emerging wasps can vary considerably among patches, depending on the relative quality of hosts and the number of females that parasitize a host. The newly revealed use of attractive and repellent pheromone compounds by C. glomerata possibly helps maximize mating success under these variable conditions.

Nocturnal Bee Pollinators Are Attracted to Guarana Flowers by Their Scents.

Floral scent is an important component of the trait repertoire of flowering plants, which is used to attract and manipulate pollinators. Despite advances during the last decades about the chemicals released by flowers, there is still a large gap in our understanding of chemical communication between flowering plants and their pollinators. We analyzed floral scents of guarana (Paullinia cupana, Sapindaceae), an economically important plant of the Amazon, using chemical analytical approaches, and determined the attractiveness of the scent to its nocturnal bee pollinators using behavioral assays in the field. Pollen loads of attracted bees were also analyzed. Inflorescences of guarana emit strong scents, both during day and at night, with some semi-quantitative differences between day- and night-time scents. Synthetic scent mixtures containing some of the identified floral scent components, including the most abundant ones, i.e., linalool and (E)-ß-ocimene, successfully attracted the nocturnal Megalopta bee pollinators. Pollen analyses revealed that many of the attracted bees had pollen grains from previous visits to guarana flowers on their bodies. Overall, our data show that guarana flowers attract nocturnal bee visitors by their strong scents and suggest that the chemical communication between this plant and its pollinators is a key component in crop production of this economically important plant species.

Incomplete synchrony of inflorescence scent and temperature patterns in Arum maculatum L. (Araceae).

In many Araceae both scent and heat production are known to temporally vary throughout anthesis, and in several species strong scents are released for pollinator attraction when thermogenesis is also strong. However, it is not known whether the temporal patterns of both scent emission and temperature are strictly synchronous and, for example, reach their maxima at the same time. We studied Arum maculatum, a brood-site deceptive species attracting its moth fly pollinators with strong fetid scents, to study temporal patterns in scent emission and temperature during anthesis. Inflorescence scents were collected and analysed by dynamic headspace and gas chromatography-mass spectrometry (GC-MS) or by proton-transfer-reaction-time-of-flight mass spectrometry (PTR-TOFMS), and the temperature of the appendix, which is the heating osmophore during pollinator attraction, was recorded by a thermocouple. We overall found that scent emission and temperature patterns were strongly correlated. However, in none of the seven studied individuals was the highest amount of scent released at times with the maximum temperature difference. Thus, patterns of scent emission and temperature are somewhat asynchronous suggesting that high scent emission rates and temporal scent patterns in plants with thermogenesis cannot be solely explained by temperature patterns. This calls for more in-depth studies to better understand the interplay between scent emission and thermogenesis.

Diacetin, a reliable cue and private communication channel in a specialized pollination system.

The interaction between floral oil secreting plants and oil-collecting bees is one of the most specialized of all pollination mutualisms. Yet, the specific stimuli used by the bees to locate their host flowers have remained elusive. This study identifies diacetin, a volatile acetylated glycerol, as a floral signal compound shared by unrelated oil plants from around the globe. Electrophysiological measurements of antennae and behavioural assays identified diacetin as the key volatile used by oil-collecting bees to locate their host flowers. Furthermore, electrophysiological measurements indicate that only oil-collecting bees are capable of detecting diacetin. The structural and obvious biosynthetic similarity between diacetin and associated floral oils make it a reliable cue for oil-collecting bees. It is easily perceived by oil bees, but can't be detected by other potential pollinators. Therefore, diacetin represents the first demonstrated private communication channel in a pollination system.

Behavioural plasticity and sex differences in host finding of a specialized bee species.

Many animals feed on flowers, and visual as well as olfactory cues are considered as most important mediators in animal-plant interactions. However, the relative importance of these cues is not well understood. Bees are the most important animal pollinators worldwide and here, we determined the importance of decoupled and combined visual and olfactory cues of Lysimachia punctata (Primulaceae) for host plant location in both sexes of the specialized, solitary bee, Macropis fulvipes (Melittidae). Lysimachia-inexperienced female bees preferred olfactory over visual cues though visual cues increased the attractiveness of olfactory ones. In experienced females, the importance of visual cues was increased. Both Lysimachia-naive and -experienced males relied more on visual cues as compared to females. This study demonstrates that the relative weighting of cues used for host plant finding depends on the sex and experience of M. fulvipes. The latter finding reveals the presence of learning-induced behavioural plasticity in host plant finding for a bee species. It may allow the bee to forage highly efficient. Visually guided female detection on flowers by males is a likely functional explanation for the differences in the weighting of visual and olfactory cues between the sexes.

Flower scent of floral oil-producing Lysimachia punctata as attractant for the oil-bee Macropis fulvipes.

Most flowers offer nectar and/or pollen as a reward for pollinators. However, some plants are known to produce mostly fatty oil in the flowers, instead of nectar. This oil is exclusively collected by specialized oil-bees, the pollinators of the oil-plants. Little is known about chemical communication in this pollination system, especially how the bees find their hosts. We collected the floral and vegetative scent emitted by oil-producing Lysimachia punctata by dynamic headspace, and identified the compounds by gas chromatography coupled to mass spectrometry. Thirty-six compounds were detected in the scent samples, several of which were flower-specific. Pentane extracts of flowers and floral oil were tested on Macropis fulvipes in a biotest. Flower and oil extracts attracted the bees, and some of the compounds identified are seldom found in the floral scent of other plants; these may have been responsible for the attraction of the bees.