oscillating their wings). Because the haltere muscles are tiny in comparison to the flight muscles, flight muscle activity completely overshadows that of the haltere muscles during flight. It is unknown how haltere muscle activity during flight differs from walking. The left and right halteres show much more variable phase relationships while walking compared with flying, which may indicate decoupling of the left and right haltere muscles. Differences between species Although halteres are always synchronized with the movements of the wings, the phase at which they oscillate differs between species. Brachyceran flies (short-antennaed) oscillate their halteres almost exactly opposite their wings (180 degrees). More ancient suborders such as the Nematecerans (long-antennaed flies), which for example include crane flies and mosquitoes, exhibit a variety of wing-haltere phasing. These observed differences in wing-haltere coordination suggest that differences in sensory neuron output also exist between species. This means that the decoding mechanisms used by the central nervous system to interpret such movements and produce adequate motor output probably also vary depending on phylogeny. Morphology The general structure of halteres are well recognized, but much variability exists between species. The more ancient families, such as tipulidae (crane flies), possess halteres with rather long stalks. This causes the haltere bulbs to be much further from the body and easily visible to the naked eye. More derived families, such as Calliphoridae (blow flies), have developed specialized structures called "calyptrae" or "squama", which are tiny flaps of wing, that cover the haltere. Pringle (1948) hypothesized that they prevented wind turbulence from affecting haltere movements, allowing more precise detection of body position, but this was never tested. The stalk of the haltere is also not always straight. Instead, the stalk's shape in more derived families tends to be reflective of the body shape of the individual. This minimizes the amount of air space between end-knobs and the sides of the abdomen and thorax. In these families, the halteres beat so close to the body that the distance between haltere and thorax is a fraction of the diameter of the haltere bulb. An extreme example of this trait is in the family Syrphidae (hoverflies), where the bulb of the haltere is positioned nearly perpendicular to the stalk. Flies typically hold their halteres at a 90-degree offset. To visualize this, if you were to imagine a person holding their arms out sideways, this would be a 180-degree offset. If that person