A permanently descended larynx is found in humans and several other species of mammals. In addition to this, the larynx of species such as fallow deer is mobile and in males it can be retracted during vocalization. The most likely explanation for the lowered retractable larynx in mammals is that it serves to exaggerate perceived body size (size exaggeration hypothesis) by decreasing the formant frequencies of calls. In this study, we quantified for the first time the elongation of the vocal tract in fallow bucks during vocalization. We also measured the effect of this vocal tract length (VTL) increase on formant frequencies (vocal tract resonances) and formant dispersion (spacing of formants). Our results show that fallow bucks increase their VTL on average by 52% during vocalization. This elongation resulted in strongly lowered formant frequencies and decreased formant dispersion. There were minimal changes to formants 1 and 2 (À0.91 and +1.9%, respectively) during vocal tract elongation, whereas formants 3, 4 and 5 decreased substantially: 18.9, 10.3 and 13.6%, respectively. Formant dispersion decreased by 12.4%. Formants are prominent in deer vocalizations and are used by males to gain information on the competitive abilities of signallers. It remains to be seen whether females also use the information that formants contain for assessing male quality before mating.

Mating success not only depends on genetic quality, but also equally on environmental factors, most prominently food availability. We investigated the interactive effects of nutritional state and body size on mating success and copula duration in yellow dung fly males (Scathophaga stercoraria; Diptera: Scathophagidae) of three body size selection lines in the laboratory in both non-competitive (single) and competitive (group) situations. Adults require protein and lipids from prey to reproduce, as well as sugars as an energy source for reproductive activity. We expected mating success to decrease with time because of sperm depletion (sugar treatment) and ⁄ or energy shortage (water treatment) relative to the control, prey plus sugar treatment. Based on physiological scaling, we also expected large-line males to become depleted either sooner because of their higher energy and sperm demands, or later because of their more efficient energy use. Average mating success indeed declined over a period of 5-7 d (or 5-15 potential copulations per male), but equally for all food treatments and body size classes. Surprisingly, water-fed and small-line males had the highest mating probability in the non-competitive setting, while in the competitive setting large-line males had the highest success. Energy-depleted males showed apparent terminal investment. Small males acquired females more readily but eventually lost them to larger males in the competitive situation. As shown before, copula duration was inversely related to body size and increased with copulation number, independent of food treatment. We conclude that sugar or prey shortage has little effect on mating success in the short term, and does not differentially affect males of different sizes.

The degree of resource monopolization relates to the distribution of resources in space and time. In general, monopolization is predicted to be high when resources (food or mates) are clumped in space, dispersed in time, and predictable in space or time. Using the yellow dung fly, Scathophaga stercoraria (Diptera: Scathophagidae), we qualitatively tested a general model that predicts the distribution of mating success among competing males based on the temporal pattern of female arrivals relative to mating time and a ranking of males in priority of access to the resource (here by body size). In a laboratory experiment approximating the natural mating situation, a constant number of males of various sizes were allowed to compete for females. As predicted, mate monopolization decreased as the temporal clumping of female arrivals increased, mediated by either a decrease in the mean or an increase in the variance of female inter-arrival times, which were manipulated independently. Males appeared to adjust their behavior to variation in female arrivals in a manner consistent with the marginal value theory of Parker and associates: forcible takeovers of females were rarer, and copula durations shorter, when females arrived regularly at short intervals. Therefore, a complex interaction of variation in intrinsic characteristics affecting male resource holding potential, mating time and stochastic, extrinsic variables increasing temporal clumping of mates generally reduces the variance in mating success among competing males and thus ultimately the opportunity and intensity of sexual selection on traits influencing male success. This theory extends operational sex ratio theory at the mechanistic, behavioral level