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The waggle dance of honey bee (Apis mellifera L.) foragers communicates to nest mates the location of a profitable food source. We used solid-phase microextraction and gas chromatography coupled with mass spectrometry to show that waggle-dancing bees produce and release two alkanes, tricosane and pentacosane, and two alkenes, Z-(9)-tricosene and Z-(9)-pentacosene, onto their abdomens and into the air. Nondancing foragers returning from the same food source produce these substances in only minute quantities. Injection of the scent significantly affects worker behavior by increasing the number of bees that exit the hive. The results of this study suggest that these compounds are semiochemicals involved in worker recruitment. By showing that honey bee waggle dancers produce and release behaviorally active chemicals, this study reveals a new dimension in the organization of honey bee foraging.

The hawkmoth Manduca sexta (Lepidoptera: Sphingidae), an experimentally favorable Lepidopteran that is highly sensitive to carbon dioxide (CO2), feeds on the nectar of a range of flowering plants, such as Datura wrightii (Solanaceae). Newly opened Datura flowers give off dramatically elevated levels of CO2 and offer ample nectar. Thus, floral CO2 emission could indicate food-source profitability. This study documents that foraging Manduca moths prefer surrogate flowers that emit high levels of CO2, characteristic of newly opened Datura flowers. We show for the first time that CO2 may play an important role in the foraging behavior of nectar-feeding insects.

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SUMMARY The tremble dance of honey bee nectar foragers is part of the communication system that regulates a colony's foraging efficiency. A forager that returns to the hive with nectar, but then experiences a long unloading delay because she has difficulty finding a nectar receiver bee, will perform a tremble dance to recruit additional nectar receiver bees. A forager that experiences a short unloading delay will perform a waggle dance to recruit more nectar foragers. A long unloading delay was until now the only known cause of tremble dancing. However, several studies suggested that factors at the food source may also cause tremble dancing. Here I test whether one of these factors, crowding of nectar foragers at the food source, stimulates tremble dancing because it causes long unloading delays. To do so, I increased the density of nectar foragers at a food source by suddenly reducing the size of an artificial feeder, and recorded the unloading delay experienced by each forager, as well as the dance she performed, if any. A forager's unloading delay was measured as the time interval between entering the hive and either (1) the first unloading contact with a nectar receiver bee, or (2) the start of the first dance, if dancing began before the first unloading contact. I also recorded the unloading delays and dances of nectar foragers that returned from natural food sources. The results show that crowding of nectar foragers at the food source increases the probability of tremble dancing, but does not cause long unloading delays, and that tremble dancers that foraged at natural food sources also often have short unloading delays. When the cause of the tremble dance is not a low supply of nectar receiver bees, the tremble dance may have a function in addition to the recruitment of nectar receiver bees.

-Two hypotheses concerning the function of the hydrocarbons emitted by waggle-dancing bees are investigated in this study. First, we test the hypothesis that the waggle-dance compounds stimulate foraging behavior. In support of this hypothesis, the number of bee departures from a hive increased as much as 46% and the number of forager visits to a feeder station increased as much as 106% when the dance compounds were introduced into the hive. Second, we test the hypothesis that the waggle-dance compounds stimulate foraging by causing experienced foragers to depart the hive to exploit known food sources. In support of this hypothesis, individually marked foragers visited an empty feeder station both more often and in greater numbers following exposure to the waggle-dance compounds than following exposure to a control substance. The results of these experiments suggest that the waggle-dance compounds function as a foraging semiochemical, operating in synergy with the waggle dance itself.waggle dance / cuticular hydrocarbon / nectar foraging / semiochemical / pheromone

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Departure of swarms from honey bee (Apis mellifera Linnaeus (Hymenoptera: Apidae)) nests is an important reproductive event for wild honey bee colonies and economically costly in managed bee colonies. The seasonal timing of swarm departure varies regionally and annually, creating challenges for honey bee management and emphasizing the potential for swarming behavior to be affected by plant-pollinator phenological mismatch. In this study, we first document variability in the timing of swarm departure across the large and heterogeneous geographical area of New Jersey over 4 years using 689 swarm-cluster observations. Second, hypothesizing that honey bee colonies adaptively tune the timing of swarm departure to match floral food-resource availability, we predicted that growing degree-days could be used to account for regional and annual variability. To test this idea, we used local weather records to determine the growing degree-day on which each swarm cluster was observed and tested for differences among climate regions and years. The state-wide mean swarm cluster date was May 15 (± 0.6 d), with moderate but significant differences among the state's five climate regions and between years. Use of degree-day information suggests that local heat accumulation can account for some climate-region differences in swarm-departure timing. Annual variation existed on a scale of only several days and was not accounted for by growing degree-days, suggesting little adaptive tuning of swarm-departure timing with respect to local heat accumulation.