Alan G. McElligott

Sexual size dimorphism in fallow deer (Dama dama): do larger, heavier males gain greater mating success?

Abstract Sexual size dimorphism may evolve as a result of both natural and sexual selection. In polygynous mammals, the main factor resulting in the evolution of large body size in males is the advantage conferred during competition for mates. In this study, we examined whether sexual selection acts on body size in mature fallow bucks (Dama dama) by examining how the following traits are inter-related: age, body (skeletal) size, body mass, prerut dominance rank, rut dominance rank and mating success. This is the first study to examine how all these factors are together related to the mating success of a large sexually dimorphic and polygynous mammal. We found that male mating success was directly related to body size, but not to body mass. However body mass was related to prerut dominance rank which was in turn strongly related to rut dominance rank, and thus there was an indirect relationship between mating success and body mass. Rut dominance rank was the variable most strongly related to mating success. Mating success among mature males was unrelated to age. We conclude that larger mature fallow bucks have advantages over other males when competing for matings, and sexual selection therefore continues to act on sexual size dimorphism in this species. Heavier fallow bucks also have advantages, but these are mediated through the dominance ranks attained by males before the rut.

Low Frequency Groans Indicate Larger and More Dominant Fallow Deer (Dama dama) Males

Background Models of honest advertisement predict that sexually selected calls should signal male quality. In most vertebrates, high quality males have larger body sizes that determine higher social status and in turn higher reproductive success. Previous research has emphasised the importance of vocal tract resonances or formant frequencies of calls as cues to body size in mammals. However, the role of the acoustic features of vocalisations as cues to other quality-related phenotypic characteristics of callers has rarely been investigated. Methodology/Principal Findings We examined whether the acoustic structure of fallow deer groans provides reliable information on the quality of the caller, by exploring the relationships between male quality (body size, dominance rank, and mating success) and the frequency components of calls (fundamental frequency, formant frequencies, and formant dispersion). We found that body size was not related to the fundamental frequency of groans, whereas larger males produced groans with lower formant frequencies and lower formant dispersion. Groans of high-ranking males were characterised by lower minimum fundamental frequencies and to a lesser extent, by lower formant dispersions. Dominance rank was the factor most strongly related to mating success, with higher-ranking males having higher mating success. The minimum fundamental frequency and the minimum formant dispersion were indirectly related to male mating success (through dominance rank). Conclusion/Significance Our study is the first to show that sexually selected vocalisations can signal social dominance in mammals other than primates, and reveals that independent acoustic components encode accurate information on different phenotypic aspects of male quality.

Age-specific survival and reproductive probabilities: evidence for senescence in male fallow deer (Dama dama)

Survival and reproduction are key features in the evolution of life–history strategies. In this study, we use capture–mark–resighting and multi–state models to examine survival senescence and reproductive senescence in six successive cohorts of fallow bucks that were studied for 16 years. We found that the overall age-specific survival probabilities of males were highly variable and the best–fitting model revealed that fallow bucks have four life–history stages: yearling, pre–reproductive, prime–age and senescent. Pre–reproductive males (2 and 3 years old) had the highest survival. Survival declined sharply after the age of 9 years, indicating that senescence had begun. When we considered reproducing and non-reproducing males separately, there was no evidence of senescence in the former, and steadily decreasing survival after the onset of social maturity in the latter. Reproduction probability also declined in older males, and thus we provide very strong evidence of senescence. Reproducers had a greater chance of reproducing again in the following year than non–reproducers. Furthermore, there were differences in the survival probabilities, with reproducers consistently surviving better than non–reproducers. In our study population, reproducers allocate more to the effort to reproduce than non–reproducers. Therefore our results indicate the generally higher phenotypic quality of reproducing males. These results, along with earlier studies on the same population, could indicate positive relationships between fitness correlates.

Mother-young recognition in an ungulate hider species: A unidirectional process

Parent‐offspring recognition is usually crucial for survival of young. In mammals, olfaction often only permits identification at short range, and vocalizations are important at longer distances. Following and hiding antipredator strategies found in newborn mammals may also affect parental recognition mechanisms. We investigated mother‐offspring recognition in fallow deer, an ungulate hider species. We analyzed the structure of adult female and fawn contact calls to determine whether they are individually distinctive and tested for mother‐offspring recognition. Only females (and not fawns) have individualized vocalizations, with the fundamental frequency as the most distinctive parameter. Playback experiments showed that fawns can distinguish the calls of their mothers from those of other females, but mothers could not discriminate their own and alien fawn calls. Thus, the vocal identification process is unidirectional. In followers, mother‐offspring acoustic recognition is mutual, and therefore the different antipredator strategies of newborn mammals may have shaped the modalities of parent‐offspring acoustic recognition.

Fallow bucks get hoarse: vocal fatigue as a possible signal to conspecifics

Many studies of sexually selected vocal communication assume that calls remain stable throughout the breeding season. However, during this period, physiological and social factors change and these can have strong effects on the structure of calls and calling rates. During the rut, fallow bucks, Dama dama, reduce their feeding and increase the time and energy spent on vocalizing and fighting to gain matings, and consequently their body condition declines greatly. The availability of matings and intensity of competition between males also change. Therefore, we predicted that male vocal signalling would vary over time in response to the changing intersexual and intrasexual selective environment. We measured the structure of fallow buck groans and the groaning rate throughout the rut. Fundamental frequency-related parameters were highest at the beginning and at the end of the rut, and lowest during the middle when most matings occur. The fundamental frequency perturbation along the groan (Jitter) remained stable throughout the rut, whereas the number of pulses and duration of the groans decreased linearly. The minimum formant dispersion did not vary significantly over the rut. Groaning rate increased towards the middle of the rut and then rapidly decreased afterwards. We suggest that changes in the structure of groans and groaning rate are associated with the declining body condition of males and variation in the availability of mating opportunities. The breakdown in some aspects of call structure towards the end of the breeding season may represent an honest signal that could be widespread in other species.

Mother goats do not forget their kids’ calls

Parent–offspring recognition is crucial for offspring survival. At long distances, this recognition is mainly based on vocalizations. Because of maturation-related changes to the structure of vocalizations, parents have to learn successive call versions produced by their offspring throughout ontogeny in order to maintain recognition. However, because of the difficulties involved in following the same individuals over years, it is not clear how long this vocal memory persists. Here, we investigated long-term vocal recognition in goats. We tested responses of mothers to their kids’ calls 7–13 months after weaning. We then compared mothers’ responses to calls of their previous kids with their responses to the same calls at five weeks postpartum. Subjects tended to respond more to their own kids at five weeks postpartum than 11–17 months later, but displayed stronger responses to their previous kids than to familiar kids from other females. Acoustic analyses showed that it is unlikely that mothers were responding to their previous kids simply because they confounded them with the new kids they were currently nursing. Therefore, our results provide evidence for strong, long-term vocal memory capacity in goats. The persistence of offspring vocal recognition beyond weaning could have important roles in kin social relationships and inbreeding avoidance.

Mating tactics of male feral goats (Capra hircus): risks and benefits

Intense competition between males for reproduction has led to the evolution of alternative mating tactics (AMTs). Feral goat males usually use a tactic called tending, in which they defend oestrous females from other males. Males may also use a second mating tactic called coursing, in which they gain access to oestrous females by disturbing a tending pair. Herein, we examine estimated mating success (EMS) and risks of using these tactics. Tending was only used by mature (≥4 years old), higher-ranking males and accounted for 75% of EMS. Coursing was used by males of all ages and dominance ranks, and accounted for 25% of EMS. Using coursing, male kids achieved 8% of EMS. Mature males achieved 92% of EMS. Both age and dominance rank were related to EMS, but age was not important after its relationship with dominance was controlled. Tending bouts were, on average, ca. 30xa0min long, while coursing bouts only averaged ca. 2xa0min. Males were more likely to suffer a butt while coursing than while tending, and formerly tending males were responsible for most butts. Kids that coursed had the highest risk of being butted. In most AMTs, there are reductions in the risks in relation to low fitness benefits. However, we found that the risks of butts during coursing were high, while our evidence suggests that the EMS was probably low. Nevertheless, the existence of an effective AMT in male feral goats may have an important influence on the intensity of sexual selection and the effective population size.