Mandibular gland secretions of meliponine worker bees: further evidence for their role in interspecific and intraspecific defence and aggression and against their role in food source signalling.

Like ants and termites some species of stingless bees (Meliponini), which are very important pollinators in the tropics, use pheromone trails to communicate the location of a food source. We present data on the communicative role of mandibular gland secretions of Meliponini that resolve a recent controversy about their importance in the laying of such trails. Volatile constituents of the mandibular glands have been erroneously thought both to elicit aggressive/defensive behaviour and to signal food source location. We studied Trigona spinipes and Scaptotrigona aff. depilis ('postica'), two sympatric species to which this hypothesis was applied. Using extracts of carefully dissected glands instead of crude cephalic extracts we analysed the substances contained in the mandibular glands of worker bees. Major components of the extracts were 2-heptanol (both species), nonanal (T. spinipes), benzaldehyde and 2-tridecanone (S. aff. depilis). The effect of mandibular gland extracts and of individual components thereof on the behaviour of worker bees near their nest and at highly profitable food sources was consistent. Independent of the amount of mandibular gland extract applied, the bees overwhelmingly reacted with defensive behaviour and were never attracted to feeders scented with mandibular gland extract or any of the synthetic chemicals tested. Both bee species are capable of using mandibular gland secretions for intra- and interspecific communication of defence and aggression and share 2-heptanol as a major pheromone compound. While confirming the role of the mandibular glands in nest defence, our experiments provide strong evidence against their role in food source signalling.

Thoracic vibrations in stingless bees (Melipona seminigra): resonances of the thorax influence vibrations associated with flight but not those associated with sound production.

Bees generate thoracic vibrations with their indirect flight muscles in various behavioural contexts. The main frequency component of non-flight vibrations, during which the wings are usually folded over the abdomen, is higher than that of thoracic vibrations that drive the wing movements for flight. So far, this has been concluded from an increase in natural frequency of the oscillating system in association with the wing adduction. In the present study, we measured the thoracic oscillations in stingless bees during stationary flight and during two types of non-flight behaviour, annoyance buzzing and forager communication, using laser vibrometry. As expected, the flight vibrations met all tested assumptions for resonant oscillations: slow build-up and decay of amplitude; increased frequency following reduction of the inertial load; and decreased frequency following an increase of the mass of the oscillating system. Resonances, however, do not play a significant role in the generation of non-flight vibrations. The strong decrease in main frequency at the end of the pulses indicates that these were driven at a frequency higher than the natural frequency of the system. Despite significant differences regarding the main frequency components and their oscillation amplitudes, the mechanism of generation is apparently similar in annoyance buzzing and forager vibrations. Both types of non-flight vibration induced oscillations of the wings and the legs in a similar way. Since these body parts transform thoracic oscillations into airborne sounds and substrate vibrations, annoyance buzzing can also be used to study mechanisms of signal generation and transmission potentially relevant in forager communication under controlled conditions.

The sound field generated by tethered stingless bees (Melipona scutellaris): inferences on its potential as a recruitment mechanism inside the hive.

In stingless bees, recruitment of hive bees to food sources involves thoracic vibrations by foragers during trophallaxis. The temporal pattern of these vibrations correlates with the sugar concentration of the collected food. One possible pathway for transferring such information to nestmates is through airborne sound. In the present study, we investigated the transformation of thoracic vibrations into air particle velocity, sound pressure, and jet airflows in the stingless bee Melipona scutellaris. Whereas particle velocity and sound pressure were found all around and above vibrating individuals, there was no evidence for a jet airflow as with honey bees. The largest particle velocities were measured 5 mm above the wings (16.0+/-4.8 mm s(-1)). Around a vibrating individual, we found maximum particle velocities of 8.6+/-3.0 mm s(-1) (horizontal particle velocity) in front of the bee's head and of 6.0+/-2.1 mm s(-1) (vertical particle velocity) behind its wings. Wing oscillations, which are mainly responsible for air particle movements in honey bees, significantly contributed to vertically oriented particle oscillations only close to the abdomen in M. scutellaris (distances < or =5 mm). Almost 80% of the hive bees attending trophallactic food transfers stayed within a range of 5 mm from the vibrating foragers. It remains to be shown, however, whether air particle velocity alone is strong enough to be detected by Johnston's organ of the bee antenna. Taking the physiological properties of the honey bee's Johnston's organ as the reference, M. scutellaris hive bees are able to detect the forager vibrations through particle movements at distances of up to 2 cm.

Spitting out information: Trigona bees deposit saliva to signal resource locations.

Stingless bees of the species Trigona spinipes (Fabricius 1793) use their saliva to lay scent trails communicating the location of profitable food sources. Extracts of the cephalic labial glands of the salivary system (not the mandibular glands, however) contain a large amount (approx. 74%) of octyl octanoate. This ester is also found on the scent-marked substrates at the feeding site. We demonstrate octyl octanoate to be a single compound pheromone which induces full trail following behaviour. The identification of the trail pheromone in this widely distributed bee makes it an ideal organism for studying the mechanism of trail following in a day flying insect.

Vibrating the food receivers: a direct way of signal transmission in stingless bees (Melipona seminigra).

An element common to the recruitment communication of eusocial bees (honey bees, stingless bees and bumble bees) are pulsed thorax vibrations generated by successful foragers within the nest. In stingless bees, foragers vibrate during the unloading of the collected food. In the present study on Melipona seminigra we demonstrate that during trophallactic contacts, the food receivers are directly vibrated by the foragers. As a consequence, both the temporal structure and the main frequency component of the forager's vibrations are directly passed on to the receiver. The vibrations are attenuated by about 17 dB on their way from the forager's thorax (velocity amplitude of the vibrations: approximately 70 mm/s) to the receiver's thorax (approximately 10 mm/s), the main amount of attenuation (about 12 dB) occurring during transmission from the head of the forager to that of the receiver. Vibrations conducted through the substrate between the forager and food receiver are comparatively small with velocity amplitudes of 0.3 mm/s. Possible ways of perception and the advantages of vibration transmission by direct contact within the recruitment context are discussed.