Key biotic and environmental constraints on the host-orientated behaviour of haematophagous Diptera are summarized. For each major group of biting Diptera, responses to host stimuli are reviewed, including activation and ranging behaviour, long-range and short-range olfactory responses and visual responses. Limitations to the comparison of results between groups of species, and the practical problems of experimental method and equipment are discussed.

Sexual recognition through wing-beat frequency matching was first demonstrated in Toxorhynchites brevipalpis, where wing-beat frequencies of males and females are similar. Here we show frequency matching in Culex quinquefasciatus, where the wing-beat frequencies of males and females differ considerably. The wing-beat frequencies converge not on the fundamental but on the nearest shared harmonic (usually female's third and male's second). Frequencies in this range are, however, too high to elicit phasic sensory-neural responses from the Johnston's organ (JO) or to drive the mosquito's motor neurons. Potential cues for frequency matching are difference tones produced by nonlinear mixing of male and female flight tones in the vibrations of the mosquito's antennae. Receptor potentials and neural-motor activity were recorded in response to difference tones produced when a mosquito was stimulated simultaneously by two tones at frequencies outside the phasic response range of the JO but within range of the antennal vibrations. We demonstrate sexual recognition through matching of flight-tone harmonics in Culex mosquitoes and suggest that difference tones are used as an error signal for frequency matching beyond the frequency range of the JO's sensory-neural range. This is the first report of acoustic distortion being exploited as a sensory cue, rather than existing as an epiphenomenon.

Anopheles gambiae, responsible for the majority of malaria deaths annually, is a complex of seven species and several chromosomal/molecular forms. The complexity of malaria epidemiology and control is due in part to An. gambiae's remarkable genetic plasticity, enabling its adaptation to a range of human-influenced habitats. This leads to rapid ecological speciation when reproductive isolation mechanisms develop [1-6]. Although reproductive isolation is essential for speciation, little is known about how it occurs in sympatric populations of incipient species [2]. We show that in such a population of "M" and "S" molecular forms, a novel mechanism of sexual recognition (male-female flight-tone matching [7-9]) also confers the capability of mate recognition, an essential precursor to assortative mating; frequency matching occurs more consistently in same-form pairs than in mixed-form pairs (p = 0.001). [corrected] Furthermore, the key to frequency matching is "difference tones" produced in the nonlinear vibrations of the antenna by the combined flight tones of a pair of mosquitoes and detected by the Johnston's organ. By altering their wing-beat frequencies to minimize these difference tones, mosquitoes can match flight-tone harmonic frequencies above their auditory range. This is the first description of close-range mating interactions in incipient An. gambiae species.