Bumble bees are of major importance, ecologically and economically as pollinators in cool and temperate biomes and as model organisms for scientific research. Chemical signals and cues have been shown to play an outstanding role in intraspecific and interspecific communication systems within and outside of a bumble bee colony. In the present review we compile and critically assess the literature on the chemical ecology of bumble bees, including cuckoo bumble bees. The development of new and more sensitive analytical tools and improvements in sociogenetic methods significantly enhanced our knowledge about chemical compounds that mediate the regulation of reproduction in the social phase of colony development, about the interactions between host bumble bees and their social parasites, about pheromones involved in mating behavior, as well as about the importance of signals, cues and context-dependent learning in foraging behavior. Our review intends to stimulate new studies on the many unresolved questions concerning the chemical ecology of these fascinating insects.

Chemical communication is ubiquitous. The identification of conserved structural elements in visual and acoustic communication is well established, but comparable information on chemical communication displays (CCDs) is lacking. We assessed the phenotypic integration of CCDs in a meta-analysis to characterize patterns of covariation in CCDs and identified functional or biosynthetically constrained modules. Poorly integrated plant CCDs (i.e. low covariation between scent compounds) support the notion that plants often utilize one or few key compounds to repel antagonists or to attract pollinators and enemies of herbivores. Animal CCDs (mostly insect pheromones) were usually more integrated than those of plants (i.e. stronger covariation), suggesting that animals communicate via fixed proportions among compounds. Both plant and animal CCDs were composed of modules, which are groups of strongly covarying compounds. Biosynthetic similarity of compounds revealed biosynthetic constraints in the covariation patterns of plant CCDs. We provide a novel perspective on chemical communication and a basis for future investigations on structural properties of CCDs. This will facilitate identifying modules and biosynthetic constraints that may affect the outcome of selection and thus provide a predictive framework for evolutionary trajectories of CCDs in plants and animals.

The pheromones used by several species of stingless bees for scent trail communication are generally assumed to be produced by the mandibular glands. Here we present strong evidence that in Trigona recursa these pheromones originate from the labial glands, which are well developed in the heads of foragers. Analysis of the behavior involved in scent marking shows that a bee extends her proboscis and rubs it over the substrate. A single scent marking event lasts for 0.59+/-0.21 s while the bee runs a stretch of 1.04+/-0.37 cm on a leaf. According to choice experiments the bees are attracted by a feeder baited with labial gland extract (84.2+/-6% of the bees choose this feeder) but repelled from a feeder baited with mandibular gland extract (only 27.5+/-13.1% of the bees choose this feeder). They do not discriminate between two clean feeders (49.6+/-3% of the bees at a feeder). 87+/-5.1% of bees already feeding leave the feeder after the application of mandibular gland extract whereas only 6.2+/-4.9% and 2.6+/-4% do so when labial gland extract or pure solvent was applied.

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-The two stingless bee species Melipona scutellaris and M. quadrifasciata recruit nestmates to a rich foraging site. We tested this with feeders up to 140 m away from the hive. Foragers of M. scutellaris communicated direction (up to 140 m) more accurately than distance (up to 30 m) whereas those of M. quadrifasciata communicated direction only up to 30 m and distance up to 40 m. Our data indicate that in both species recruitment is divided into two temporal phases. Whereas in an initial phase alarmed nestmates search for food at random, bees leaving the hive in the following phase are obviously provided with information about its specific location. As a consequence after 35 minutes (M. scutellaris) and 85 minutes (M. quadrifasciata), respectively, significantly more newcomers arrive at the feeder than at an identical control feeder. The differences found in the recruitment success of M. scutellaris and M. quadrifasciata are discussed in regard to the different demands of their natural habitats.

-Some species of stingless bees of the genus Melipona were reported to scent mark food sources but little is known about the chemical signals involved. We studied the origin and some properties of such scent marks in M. seminigra. Results from choice experiments suggested that the bees do not scent mark the food (sugar water) itself and that abdominal droplets were excluded as the signal source. Extracts of the most distal tarsomeres, however, attracted recruits in the same way as natural scent marks. We conclude that M. seminigra scent marks a food source by leaving "footprints" secreted at the leg tips. The footprints of at least 40 visits were needed to effectively scent mark. The chemical signal has an active range of about 1 m and its effect persists for about 2 hours. In the absence of footprints no scent marking effect was seen. This finding excludes the importance of mandibular gland secretions (if at all present) for the scent marking observed.stingless bee / Melipona / scent marking / footprint substance / recruitment

We examined the ability of stingless bees to recruit nest mates to a food source (i) in group foraging species laying pheromone trails from the food to the nest (Trigona recursa SMITH, T. hypogea SILVESTRI, Scaptotrigona depilis MOURE), (ii) in solitary foraging species with possible but still doubtful communication of food location inside the nest (Melipona seminigra FRIESE, M. favosa orbignyi GUÉRIN), and (iii) in species with a less precise (Nannotrigona testaceicornis LEP., Tetragona clavipes FAB.) or no communication (Frieseomelitta varia LEP.). The bees were allowed to collect food (sugar solution or liver in the necrophageous species) ad libitum and the forager number to accumulate, as it would do under normal unrestrained conditions. The median number of bees collecting differed considerably among the species (1.0 -1436.5). It was highest in the species employing scent trails. The time course of recruitment was characteristic for most of the species and largely independent of the number of foragers involved. The two Melipona species recruited other bees significantly faster than T. recursa, S. depilis, and N. testaceicornis during the first 10 to 30 minutes of an experiment. In species laying a scent trail to guide nestmates to a food source the first recruits appeared with a delay of several minutes followed by a quick increase in forager number. The median time required to recruit all foragers available differed among the species between 95.0 and 240.0 min. These differences can at least partly be explained by differences in the recruitment mechanisms and do not simply follow from differences in colony biomass.

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.