Megan T. Wyman

Amplitude of bison bellows reflects male quality, physical condition and motivation

Sound amplitude (measured as sound pressure level) is an acoustical parameter that has received little attention within communication research, especially in mammals. Although difficult to measure in the field, amplitude is a potentially important parameter of sexually selected signals. In North American plains bison, Bison bison, ‘bellows’ are low, guttural vocalizations made by bulls during the breeding season in the context of male–male contests. It has been hypothesized that bison use bellow amplitude to assess males during male–male competition or female mate choice. In this study, we tested the hypothesis that amplitude is significantly related to measures of bull competitive ability (quality, condition and motivation), and thus, could function as a sexually selected signal. During peak rut over 2 years, courtship and threat behaviours were recorded daily. During observation sessions, bellow amplitude was measured as peak sound pressure levels using a sound level meter. Subsequent genetic parentage analysis determined offspring sired by males. Based on aspects of signalling and game theory, we predicted positive associations between amplitude and mating and reproductive success, dominance, physical condition, motivation to retain females, age and weight. Our results supported a positive association between amplitude and both physical condition and motivation. Conversely, the results showed a negative association between amplitude and quality, as measured by mating and reproductive success. Supporting evidence and alternative hypotheses for these results are explored. Our findings provide support for the notion that bellow amplitude could be used as a sexually selected signal to assess rival males during male–male competition.

Acoustic cues to size and quality in the vocalizations of male North American bison, Bison bison

Source-filter theory provides a framework to interpret the acoustic structure of vertebrate vocalizations in relation to biophysical production, and it predicts that specific acoustic parameters can encode information about callers. Because formant frequencies are determined by vocal tract dimensions, with longer vocal tracts producing lower formants, they can be reliable indicators of body size, as well as other important traits. In polygynous species, reliable acoustic cues to fitness-related traits are expected to be under strong sexual selection pressure through male competition and/or female choice. This study investigates whether formant frequencies of male North American bison bellow vocalizations encode information about fitness-related caller attributes. Bison exhibit male-dominance female-defence polygyny, with dominance displays involving bellows. We hypothesized that physical attributes (mass, age) would predict formants and that formants would in turn predict quality indices (dominance, copulations, offspring sired). Our results showed that heavier bulls produced lower formants and that lower formants predicted higher mating success (copulations), even when controlling for mass. Given positive associations between mating success, dominance and reproductive success (offspring sired) in bison, we conclude that bellows with lower formants reflect greater fitness in bulls. We discuss the importance of reliable acoustic cues to size and quality indices in sexual selection contexts.

Visualization of system dynamics using phasegrams

A new tool for visualization and analysis of system dynamics is introduced: the phasegram. Its application is illustrated with both classical nonlinear systems (logistic map and Lorenz system) and with biological voice signals. Phasegrams combine the advantages of sliding-window analysis (such as the spectrogram) with well-established visualization techniques from the domain of nonlinear dynamics. In a phasegram, time is mapped onto the x-axis, and various vibratory regimes, such as periodic oscillation, subharmonics or chaos, are identified within the generated graph by the number and stability of horizontal lines. A phasegram can be interpreted as a bifurcation diagram in time. In contrast to other analysis techniques, it can be automatically constructed from time-series data alone: no additional system parameter needs to be known. Phasegrams show great potential for signal classification and can act as the quantitative basis for further analysis of oscillating systems in many scientific fields, such as physics (particularly acoustics), biology or medicine.

A search for principles of disability using experimental impairment of Drosophila melanogaster

The results of life table experiments to determine the effects of artificial impairment (leg amputation) in 7500 Drosophila melanogaster adults revealed that the extent to which life expectancy was reduced in impaired individuals was conditional on: (1) leg location and number amputated--front leg had greatest impact and the number of legs amputated directly correlated with mortality impact; (2) age of amputation--the greatest relative reduction in remaining life expectancy occurred when young flies were impaired; (3) vial orientation--mortality in impaired flies was the least when vials held upside-down (most friendly environment) and the greatest when they were right-side up (least friendly environment); and (4) sex--male mortality was increased more than female mortality in nearly all impairment treatments. These results were used to formulate a set of general principles of disability that would apply not only to humans but to all organisms.

Variability of Female Responses to Conspecific vs. Heterospecific Male Mating Calls in Polygynous Deer: An Open Door to Hybridization?

Males of all polygynous deer species (Cervinae) give conspicuous calls during the reproductive season. The extreme interspecific diversity that characterizes these vocalizations suggests that they play a strong role in species discrimination. However, interbreeding between several species of Cervinae indicates permeable interspecific reproductive barriers. This study examines the contribution of vocal behavior to female species discrimination and mating preferences in two closely related polygynous deer species known to hybridize in the wild after introductions. Specifically, we investigate the reaction of estrous female red deer (Cervus elaphus) to playbacks of red deer vs. sika deer (Cervus nippon) male mating calls, with the prediction that females will prefer conspecific calls. While on average female red deer preferred male red deer roars, two out of twenty females spent more time in close proximity to the speaker broadcasting male sika deer moans. We suggest that this absence of strict vocal preference for species-specific mating calls may contribute to the permeability of pre-zygotic reproductive barriers observed between these species. Our results also highlight the importance of examining inter-individual variation when studying the role of female preferences in species discrimination and intraspecific mate selection.

Evidence of biphonation and source–filter interactions in the bugles of male North American wapiti (Cervus canadensis)

ABSTRACT With an average male body mass of 320 kg, the wapiti, Cervus canadensis, is the largest extant species of Old World deer (Cervinae). Despite this large body size, male wapiti produce whistle-like sexual calls called bugles characterised by an extremely high fundamental frequency. Investigations of the biometry and physiology of the male wapitis relatively large larynx have so far failed to account for the production of such a high fundamental frequency. Our examination of spectrograms of male bugles suggested that the complex harmonic structure is best explained by a dual-source model (biphonation), with one source oscillating at a mean of 145 Hz (F0) and the other oscillating independently at an average of 1426 Hz (G0). A combination of anatomical investigations and acoustical modelling indicated that the F0 of male bugles is consistent with the vocal fold dimensions reported in this species, whereas the secondary, much higher source at G0 is more consistent with an aerodynamic whistle produced as air flows rapidly through a narrow supraglottic constriction. We also report a possible interaction between the higher frequency G0 and vocal tract resonances, as G0 transiently locks onto individual formants as the vocal tract is extended. We speculate that male wapiti have evolved such a dual-source phonation to advertise body size at close range (with a relatively low-frequency F0 providing a dense spectrum to highlight size-related information contained in formants) while simultaneously advertising their presence over greater distances using the very high-amplitude G0 whistle component. Highlighted Article: North American wapitis produce extremely high-pitched bugles that are incompatible with the dimensions of their vocal folds. Anatomical and acoustic investigations suggest plausible mechanisms responsible for the production of this extraordinary vocalisation.

Response of red deer stags (Cervus elaphus) to playback of harsh versus common roars

Red deer stags (Cervus elaphus) give two distinct types of roars during the breeding season, the “common roar” and the “harsh roar.” Harsh roars are more frequent during contexts of intense competition, and characterized by a set of features that increase their perceptual salience, suggesting that they signal heightened arousal. While common roars have been shown to encode size information and mediate both male competition and female choice, to our knowledge, the specific function of harsh roars during male competition has not yet been studied. Here, we investigate the hypothesis that the specific structure of male harsh roars signals high arousal to competitors. We contrast the behavioral responses of free ranging, harem-holding stags to the playback of harsh roars from an unfamiliar competitor with their response to the playback of common roars from the same animal. We show that males react less strongly to sequences of harsh roars than to sequences of common roars, possibly because they are reluctant to escalate conflicts with highly motivated and threatening unfamiliar males in the absence of visual information. While future work should investigate the response of stags to harsh roars from familiar opponents, our observations remain consistent with the hypothesis that harsh roars may signal motivation during male competition, and illustrate how intrasexual selection can contribute to the diversification of male vocal signals.

Interactive patterns of vocal communication in African elephant herds (Loxodonta africana)

This study examines the interactive nature of vocalizations produced within African elephant herds during waterhole visits at Etosha National Park, Namibia. The temporal organization and physical characteristics of 1025 vocalizations, documented within 14.8 h of acoustic field recordings, were analyzed using a variety of statistical tests. Temporal distribution analyses indicated that calls were clumped in time (Kologorov–Smirnov, average significant p=0.001 58) and the majority of the calls occurred after herds began the process of departing a waterhole (Wilcoxon signed rank, p=0.0012). In addition, 84% of all calls began within 30 s of another call. Based on the analysis of the temporal patterning of calls, a conversational series of vocalizations was defined as calls separated by less than 30 s of silence. The majority of calls in a conversational series are overlapping or contiguous, indicating that multiple elephants are involved in the exchange. A variety of physical characteristics (duration, fr...

Vocal tract modelling in fallow deer: are male groans nasalized?

ABSTRACT Males of several species of deer have a descended and mobile larynx, resulting in an unusually long vocal tract, which can be further extended by lowering the larynx during call production. Formant frequencies are lowered as the vocal tract is extended, as predicted when approximating the vocal tract as a uniform quarter wavelength resonator. However, formant frequencies in polygynous deer follow uneven distribution patterns, indicating that the vocal tract configuration may in fact be rather complex. We CT-scanned the head and neck region of two adult male fallow deer specimens with artificially extended vocal tracts and measured the cross-sectional areas of the supra-laryngeal vocal tract along the oral and nasal tracts. The CT data were then used to predict the resonances produced by three possible configurations, including the oral vocal tract only, the nasal vocal tract only, or combining the two. We found that the area functions from the combined oral and nasal vocal tracts produced resonances more closely matching the formant pattern and scaling observed in fallow deer groans than those predicted by the area functions of the oral vocal tract only or of the nasal vocal tract only. This indicates that the nasal and oral vocal tracts are both simultaneously involved in the production of a non-human mammal vocalization, and suggests that the potential for nasalization in putative oral loud calls should be carefully considered. Highlighted Article: CT scans of fallow deer specimens show that the nasal cavities are likely to be involved in the production of their groans.

Introgression of exotic Cervus (nippon and canadensis) into red deer (Cervus elaphus) populations in Scotland and the English Lake District

Abstract Since the mid‐19th century, multiple introductions of Japanese sika deer (Cervus nippon nippon) and North American wapiti (C. canadensis) have taken place in the British Isles. While wapiti have generally been unsuccessful, sika have been very successful, especially in Scotland where they now overlap at least 40% of the range of native red deer (C. elaphus). Hybridization between these two species and red deer has been demonstrated in captivity and in the wild. Using a panel of 22 microsatellite loci that are highly diagnostic between red deer and sika, and moderately diagnostic between red deer and wapiti, we investigated the extent of introgression between these species in 2,943 deer sampled from around Scotland and from the English Lake District using the Bayesian clustering software STRUCTURE. We also used a diagnostic mitochondrial marker for red deer and sika. Our survey extends previous studies indicating little introgression of wapiti nuclear alleles into red deer, in particular in Northern Scotland, Kintyre, and the Lake District. We found a new area of extensive sika introgression in South Kintyre. In the North Highlands, we show for the first time geographically scattered evidence of past hybridization followed by extensive backcrossing, including one red‐like individual with sika introgression, two sika‐like individuals with red deer introgression, and six individuals that were apparently pure sika at the nuclear markers assessed but which carried red deer mitochondria. However, there has not been a collapse of assortative mating in this region. Similarly, in the English Lake District red deer, we found only traces of past sika introgression. No sika alleles were detected in the Central Highlands or the Hebridean red deer refugia. We make suggestions for management to prevent further spread of sika alleles into red deer and vice versa.