What is the guiding behavior?                               

The reports of native honeygatherers, and the research done on those claims, have shown that the guiding honeyguides utilize specific behaviors in order to successfully lead another organsim to a bee colony. The guiding has been known to take place regardless of season, by both males and females, and at any time of the day. This has been researched in regards to their interactions with humans, although the tribes themselves claim that people learned from watching the honeyguides guiding a ratel. Ratels, or honey badgers, have thick fur that would be a good defense against the bee stings. (Image: Walter A. Weber)

There are tribal tattoos and fables that indicate that the tribes were aware of this behavior for centuries. In the modern day, honeyguides will even follow motor vehicles and boats, and can be attracted by the sound of chopping wood or smoke. Greater Honeyguides persistently attempt to guide nonresponding humans, suggesting it is the humans that provide the greatest stimulus for this behavior (Dean et. al.).

Ratels are traditionally thought to be the original organisms that helped this behavior develop in honeyguides. Some scientists believe that this relationship is unlikely. Greater Honeyguides have been observed exhibiting guiding behavior in the presence of mongoose and baboon species, and neither responded. This is believed to indicate that the behavior may be released by the presence of non-symbiont mammals, including honey badgers. The distribution of the honeyguide and the honey badger overlap only in eastern and southern Africa. Greater Honeyguides do not live in the arid southwest and the tropical rainforest, where the honey badger does. The honey badger has poor hearing, making it hard to hear a guiding call, and two honey badgers who were exposed to recordings of a Greater Honeyguide chattering vocalization showed no recognition or response to the call. They are also nocturnal and have poor eyesight and climbing ability. (Dean et. al.) However, many people still refer to the ratel-honeyguide interaction as the initial example of the guiding behavior. Until a honeyguide is observed guiding a ratel in the wild, it cannot be proven.

Honeyguides have been observed visiting bee colonies repeatedly, including ones that are inaccessible by them. If it was a time when the bees were docile, they might even fly to the entrance of the nest and stick their head in. The information that the birds gather during these visits allows them to use their goal-oriented guiding behavior. They change their calls and their flight pattern in order to indicate for another organism to follow them. They fly directly to a nest that they know, returning constantly and at shorter intervals to "check up" on the followers. They begin using lower and lower perches, which may not be an indication of the proximity of the nest but insteads a result of the bird becoming less and less frightened of the honey-gatherers.

The decreasing distance between stops may represent an "area-restricted search" that many animals perform when close to their goal, be it food, hosts, or homes (Isack and Reyer). The animal spends more time scanning the area and is then likely to detect signs of the goal, such as seeing or hearing bees, and then  are able to make any directional corrections necessary to avoid missing the nest. It may be more economical for the bird to use longer stopping distances early in the search in order to limit the amount of energy-spending takeoffs and landings.  

There are seventeen species of honeyguide,  but only two use deliberate guiding behavior - the greater (black-throated) honeyguide, and the the scaly-throated (variegated) honeyguide. All of the species eat wax and are brood parasites.  All are tough and aggressive birds, as they harass brood host birds, mob wax sources, or lead humans. Honeyguides tend to be solitary, but dozens of birds of up to four species can converge upon one wax site. Scaly-throated honeyguides will push aside greater and lesser honeyguides, but immature greater honeyguides out-bully all other species.

Several of the African honeyguides, such as the greater and scaly-throated honeyguides, consistently follow and watch human activities. They fly around campsites and inspect everything from tents to cars.  These guiding honeyguides also routinely inspect colonies that are inaccessible to them, which gives them the ability to immediately lead other organisms to their location when presented with the opportunity.