Grooming and pollen manipulation in bees (Apoidea): the nature and evolution of movements involving the foreleg

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Abstract
Three modes of self cleaning occur in insects: nibbling by the maxillae, scraping one structure by another in one direction only, and rubbing back and forth while the respective parts are in continuous contact. This paper describes a comprehensive and comparative account of this behaviour in bees, with special reference to the cleaning of or by the forelegs.Bees, like all Hymenoptera, clean various parts of the head, including the mouthparts and the antennae, with the forelegs. Lower Hymenoptera scrape each antenna with either foreleg; in the species of Aculeata that possess the antenna cleaner (strigil) on the foreleg, only the ipsilateral leg is used. The thoracic dorsum of most bees, as in many sphecoid wasps, is scraped in a forward direction by the middle leg;Triepeolusspp., however, use the hind leg, and the Anthophorinae the foreleg.Some beetles and lacewings clean their forelegs in the mouthparts by nibbling and scraping. Most higher Hymenoptera as a rule scrape the foreleg between the ipsilateral maxilla and the labium; bees, however, clamp the foreleg between the flexed ipsilateral middle leg and then scrape it. An evolution of this behaviour is postulated via several intermediate forms derived from original stepping movements.Halictidae and Andrenidae clamp the foreleg for scraping underneath the middle tibia, whereas all other bees nearly always clamp it underneath the middle basitarsus. Very similar movements are used in various species for transferring pollen, oil, or nest materials from the foreleg to the middle leg.It is argued that the original way of pollen carrying in bees must have been by filling the crop through direct eating or by scraping pollen off the foreleg between the ipsilateral maxilla and the labium. The latter movement is widespread among bees and is homologous to the normal foreleg cleaning in the mouthparts of most other Hymenoptera. The efficiency of this behaviour is enhanced in many lower bees by a comb on the galea, which is the homologue of a similar structure widespread among aculeate wasps. In higher bees, Apidae and Anthophoridae, the galeal comb is replaced by an equifunctional stipes comb. Many bees have neither of these types of maxillary combs.