Holding tight on feathers - structural specializations and attachment properties of the avian ectoparasite Crataerina pallida (Diptera, Hippoboscidae)

The louse fly is an obligate blood-sucking ecto-parasite of the common swift Due to reduction of the wings, is unable to fly, thus an effective and reliable attachment to their host's plumage is of outmost importance. Its attachment system shows several modifications in comparison to other calyptrate flies. The most prominent ones are the large tridentate claws and the dichotomously shaped setae located on the pulvilli. Based on data from morphological analysis, confocal laser scanning microscopy, cryo-scanning electron microscopy and traction force experiments, performed on native (feathers) as well as artificial substrates (glass, epoxy-resin and silicone rubber), we showed that the entire attachment system is highly adapted to the fly's lifestyle as an ectoparasite. The claws in particular are the main contributor to strong attachment to the host. Resulting attachment forces on feathers make it impossible to detach without damage of feathers or legs of the fly itself. Well-developed pulvilli are responsible for the attachment to smooth surfaces. Both dichotomously shaped setae and high setal density explain high safety factors observed on smooth substrates. For the first time, we demonstrated a material gradient within the setae with soft, resilin dominated apical tips and stiff, more sclerotized bases in Diptera. The empodium seems not to be directly involved in the attachment process, but it might operate as a cleaning device and may be essential to maintain the functionality of the entire attachment system.