Roars, groans and moans: Anatomical correlates of vocal diversity in polygynous deer.

Eurasian deer are characterized by the extraordinary diversity of their vocal repertoires. Male sexual calls range from roars with relatively low fundamental frequency (hereafter fo ) in red deer Cervus elaphus, to moans with extremely high fo in sika deer Cervus nippon, and almost infrasonic groans with exceptionally low fo in fallow deer Dama dama. Moreover, while both red and fallow males are capable of lowering their formant frequencies during their calls, sika males appear to lack this ability. Female contact calls are also characterized by relatively less pronounced, yet strong interspecific differences. The aim of this study is to examine the anatomical bases of these inter-specific and inter-sexual differences by identifying if the acoustic variation is reflected in corresponding anatomical variation. To do this, we investigated the vocal anatomy of male and female specimens of each of these three species. Across species and sexes, we find that the observed acoustic variability is indeed related to expected corresponding anatomical differences, based on the source-filter theory of vocal production. At the source level, low fo is associated with larger vocal folds, whereas high fo is associated with smaller vocal folds: sika deer have the smallest vocal folds and male fallow deer the largest. Red and sika deer vocal folds do not appear to be sexually dimorphic, while fallow deer exhibit strong sexual dimorphism (after correcting for body size differences). At the filter level, the variability in formants is related to the configuration of the vocal tract: in fallow and red deer, both sexes have evolved a permanently descended larynx (with a resting position of the larynx much lower in males than in females). Both sexes also have the potential for momentary, call-synchronous vocal tract elongation, again more pronounced in males than in females. In contrast, the resting position of the larynx is high in both sexes of sika deer and the potential for further active vocal tract elongation is virtually absent in both sexes. Anatomical evidence suggests an evolutionary reversal in larynx position within sika deer, that is, a secondary larynx ascent. Together, our observations confirm that the observed diversity of vocal behaviour in polygynous deer is supported by strong anatomical differences, highlighting the importance of anatomical specializations in shaping mammalian vocal repertoires. Sexual selection is discussed as a potential evolutionary driver of the observed vocal diversity and sexual dimorphisms.

Rutting vocal display in male impala (Aepyceros melampus) and overlap with alarm context.

BACKGROUND: The rutting vocal display of male impala Aepyceros melampus is unique for its complexity among ruminants. This study investigates bouts of rutting calls produced towards potential mates and rival males by free-ranging male impala in Namibia. In particular, a comparison of male rutting and alarm snorts is conducted, inspired by earlier findings of mate guarding by using alarm snorts in male topi Damaliscus lunatus. RESULTS: Rutting male impala produced 4-38 (13.5 ± 6.5) rutting calls per bout. We analyzed 201 bouts, containing in total 2709 rutting calls of five types: continuous roars produced within a single exhalation-inhalation cycle; interrupted roars including few exhalation-inhalation cycles; pant-roars distinctive by a pant-phase with rapidly alternating inhalations and exhalations; usual snorts lacking any roar part; and roar-snorts starting with a short roar part. Bouts mostly started and ended with usual snorts. Continuous roars were the shortest roars. The average duration of the exhalatory phase was longest in the continuous roars and shortest in the pant-roars. The average fundamental frequency (49.7-51.4 Hz) did not differ between roar types. Vocal tract length, calculated by using measurements of the first four vocal tract resonances (formants), ranged within 381-382 mm in all roar types. In the studied male impala, rutting snorts within bouts of rutting calls were longer and had higher values of the upper quartile in the call spectra than alarm snorts produced towards potential danger. CONCLUSIONS: Additional inhalations during the emission of the interrupted and pant-roars prolong their duration compared to the continuous roars but do not affect the fundamental frequency or the degree of larynx retraction while roaring. Alarm snorts are separated from one another by large intervals, whereas the intervals between rutting snorts within bouts are short. Sometimes, rutting snorts alternate with roars, whereas alarm snorts do not. Therefore, it is not the acoustic structure of individual snorts but the temporal sequence and the occasional association with another call type that defines snorts as either rutting or alarm snorts. The rutting snorts of male impala may function to attract the attention of receptive females and delay their departure from a male's harem or territory.

Savannah roars: The vocal anatomy and the impressive rutting calls of male impala (Aepyceros melampus) - highlighting the acoustic correlates of a mobile larynx.

A retractable larynx and adaptations of the vocal folds in the males of several polygynous ruminants serve for the production of rutting calls that acoustically announce larger than actual body size to both rival males and potential female mates. Here, such features of the vocal tract and of the sound source are documented in another species. We investigated the vocal anatomy and laryngeal mobility including its acoustical effects during the rutting vocal display of free-ranging male impala (Aepyceros melampus melampus) in Namibia. Male impala produced bouts of rutting calls (consisting of oral roars and interspersed explosive nasal snorts) in a low-stretch posture while guarding a rutting territory or harem. For the duration of the roars, male impala retracted the larynx from its high resting position to a low mid-neck position involving an extensible pharynx and a resilient connection between the hyoid apparatus and the larynx. Maximal larynx retraction was 108 mm based on estimates in video single frames. This was in good concordance with 91-mm vocal tract elongation calculated on the basis of differences in formant dispersion between roar portions produced with the larynx still ascended and those produced with maximally retracted larynx. Judged by their morphological traits, the larynx-retracting muscles of male impala are homologous to those of other larynx-retracting ruminants. In contrast, the large and massive vocal keels are evolutionary novelties arising by fusion and linear arrangement of the arytenoid cartilage and the canonical vocal fold. These bulky and histologically complex vocal keels produced a low fundamental frequency of 50 Hz. Impala is another ruminant species in which the males are capable of larynx retraction. In addition, male impala vocal folds are spectacularly specialized compared with domestic bovids, allowing the production of impressive, low-frequency roaring vocalizations as a significant part of their rutting behaviour. Our study expands knowledge on the evolutionary variation of vocal fold morphology in mammals, suggesting that the structure of the mammalian sound source is not always human-like and should be considered in acoustic analysis and modelling.

The remarkable vocal anatomy of the koala (Phascolarctos cinereus): insights into low-frequency sound production in a marsupial species.

Koalas are characterised by a highly unusual vocal anatomy, with a descended larynx and velar vocal folds, allowing them to produce calls at disproportionately low frequencies. Here we use advanced imaging techniques, histological data, classical macroscopic dissection and behavioural observations to provide the first detailed description and interpretation of male and female koala vocal anatomy. We show that both males and females have an elongated pharynx and soft palate, resulting in a permanently descended larynx. In addition, the hyoid apparatus has a human-like configuration in which paired dorsal, resilient ligaments suspend the hyoid apparatus from the skull, while the ventral parts tightly connect to the descended larynx. We also show that koalas can retract the larynx down into the thoracic inlet, facilitated by a dramatic evolutionary transformation of the ventral neck muscles. First, the usual retractors of the larynx and the hyoid have their origins deep in the thorax. Secondly, three hyoid muscles have lost their connection to the hyoid skeleton. Thirdly, the genioglossus and geniohyoid muscles are greatly increased in length. Finally, the digastric, omohyoid and sternohyoid muscles, connected by a common tendinous intersection, form a guiding channel for the dynamic down-and-up movements of the ventral hyoid parts and the larynx. We suggest that these features evolved to accommodate the low resting position of the larynx and assist in its retraction during call production. We also confirm that the edges of the intra-pharyngeal ostium have specialised to form the novel, extra-laryngeal velar vocal folds, which are much larger than the true intra-laryngeal vocal folds in both sexes, but more developed and specialised for low frequency sound production in males than in females. Our findings illustrate that strong selection pressures on acoustic signalling not only lead to the specialisation of existing vocal organs but can also result in the evolution of novel vocal structures in both sexes.

Altai pika (Ochotona alpina) alarm calls: individual acoustic variation and the phenomenon of call-synchronous ear folding behavior.

Non-hibernating pikas collect winter food reserves and store them in hay piles. Individualization of alarm calls might allow discrimination between colony members and conspecifics trying to steal food items from a colony pile. We investigated vocal posture, vocal tract length, and individual acoustic variation of alarm calls, emitted by wild-living Altai pikas Ochotona alpina toward a researcher. Recording started when a pika started calling and lasted as long as possible. The alarm call series of 442 individual callers from different colonies consisted of discrete short (0.073-0.157 s), high-frequency (7.31-15.46 kHz), and frequency-modulated calls separated by irregular intervals. Analysis of 442 discrete calls, the second of each series, revealed that 44.34% calls lacked nonlinear phenomena, in 7.02% nonlinear phenomena covered less than half of call duration, and in 48.64% nonlinear phenomena covered more than half of call duration. Peak frequencies varied among individuals but always fitted one of three maxima corresponding to the vocal tract resonance frequencies (formants) calculated for an estimated 45-mm oral vocal tract. Discriminant analysis using variables of 8 calls per series of 36 different callers, each from a different colony, correctly assigned over 90% of the calls to individuals. Consequently, Altai pika alarm calls are individualistic and nonlinear phenomena might further increase this acoustic individualization. Additionally, video analysis revealed a call-synchronous, very fast (0.13-0.23 s) folding, depression, and subsequent re-expansion of the pinna confirming an earlier report of this behavior that apparently contributes to protecting the hearing apparatus from damage by the self-generated high-intensity alarm calls.

Unusually high-pitched neonate distress calls of the open-habitat Mongolian gazelle (Procapra gutturosa) and their anatomical and hormonal predictors.

In neonate ruminants, the acoustic structure of vocalizations may depend on sex, vocal anatomy, hormonal profiles and body mass and on environmental factors. In neonate wild-living Mongolian gazelles Procapra gutturosa, hand-captured during biomedical monitoring in the Daurian steppes at the Russian-Mongolian border, we spectrographically analysed distress calls and measured body mass of 22 individuals (6 males, 16 females). For 20 (5 male, 15 female) of these individuals, serum testosterone levels were also analysed. In addition, we measured relevant dimensions of the vocal apparatus (larynx, vocal folds, vocal tract) in one stillborn male Mongolian gazelle specimen. Neonate distress calls of either sex were high in maximum fundamental frequency (800-900 Hz), but the beginning and minimum fundamental frequencies were significantly lower in males than in females. Body mass was larger in males than in females. The levels of serum testosterone were marginally higher in males. No correlations were found between either body mass or serum testosterone values and any acoustic variable for males and females analysed together or separately. We discuss that the high-frequency calls of neonate Mongolian gazelles are more typical for closed-habitat neonate ruminants, whereas other open-habitat neonate ruminants (goitred gazelle Gazella subgutturosa, saiga antelope Saiga tatarica and reindeer Rangifer tarandus) produce low-frequency (<200 Hz) distress calls. Proximate cause for the high fundamental frequency of distress calls of neonate Mongolian gazelles is their very short, atypical vocal folds (4 mm) compared to the 7-mm vocal folds of neonate goitred gazelles, producing distress calls as low as 120 Hz.

Remarkable vocal identity in wild-living mother and neonate saiga antelopes: a specialization for breeding in huge aggregations?

Saiga antelopes Saiga tatarica tatarica give birth in large aggregations, and offspring follow the herd soon after birth. Herding is advantageous as anti-predator strategy; however, communication between mothers and neonates is strongly complicated in large aggregations. Individual series of nasal and oral contact calls of mother and neonate saiga antelopes were selected from recordings made with automated recording systems placed near the hiding neonates on the saiga breeding grounds in Northern Kazakhstan during synchronized parturitions of 30,000 calving females. We used for comparison of the acoustic structure of nasal and oral contact calls 168 nasal calls of 18 mothers, 192 oral calls of 21 mothers, 78 nasal calls of 16 neonates, and 197 oral calls of 22 neonates. In the oral calls of either mothers or neonates, formant frequencies were higher and the duration was longer than in the nasal calls, whereas fundamental frequencies did not differ between oral and nasal calls. Discriminant function analysis (DFA) based on six acoustic variables, accurately classified individual identity for 99.4% of oral calls of 18 mothers, for 89.3% of nasal calls of 18 mothers, and for 94.4% of oral calls of 18 neonates. The average value of correct classification to individual was higher in mother oral than in mother nasal calls and in mother oral calls than in neonate oral calls; no significant difference was observed between mother nasal and neonate oral calls. Variables mainly responsible for vocal identity were the fundamental frequency and the second and third formants in either mothers or neonates, and in either nasal or oral calls. The high vocal identity of mothers and neonates suggests a powerful potential for the mutual mother-offspring recognition in dense aggregations of saiga antelopes as an important component of their survival strategy.

Complex vibratory patterns in an elephant larynx.

Elephants' low-frequency vocalizations are produced by flow-induced self-sustaining oscillations of laryngeal tissue. To date, little is known in detail about the vibratory phenomena in the elephant larynx. Here, we provide a first descriptive report of the complex oscillatory features found in the excised larynx of a 25 year old female African elephant (Loxodonta africana), the largest animal sound generator ever studied experimentally. Sound production was documented with high-speed video, acoustic measurements, air flow and sound pressure level recordings. The anatomy of the larynx was studied with computed tomography (CT) and dissections. Elephant CT vocal anatomy data were further compared with the anatomy of an adult human male. We observed numerous unusual phenomena, not typically reported in human vocal fold vibrations. Phase delays along both the inferior-superior and anterior-posterior (A-P) dimension were commonly observed, as well as transverse travelling wave patterns along the A-P dimension, previously not documented in the literature. Acoustic energy was mainly created during the instant of glottal opening. The vestibular folds, when adducted, participated in tissue vibration, effectively increasing the generated sound pressure level by 12 dB. The complexity of the observed phenomena is partly attributed to the distinct laryngeal anatomy of the elephant larynx, which is not simply a large-scale version of its human counterpart. Travelling waves may be facilitated by low fundamental frequencies and increased vocal fold tension. A travelling wave model is proposed, to account for three types of phenomena: A-P travelling waves, 'conventional' standing wave patterns, and irregular vocal fold vibration.

Vocal anatomy, tongue protrusion behaviour and the acoustics of rutting roars in free-ranging Iberian red deer stags (Cervus elaphus hispanicus).

Roaring in rutting Iberian red deer stags Cervus elaphus hispanicus is unusual compared to other subspecies of red deer, which radiated from the Iberian refugium after the last glacial maximum. In all red deer stags, the larynx occupies a permanent low mid-neck resting position and is momentarily retracted almost down to the rostral end of the sternum during the production of rutting calls. Simultaneous with the retraction of the larynx, male Iberian red deer pronouncedly protrude the tongue during most of their rutting roars. This poses a mechanical challenge for the vocal tract (vt) and for the hyoid apparatus, as tongue and larynx are strongly pulled in opposite directions. This study (i) examines the vocal anatomy and the acoustics of the rutting roars in free-ranging male C. e. hispanicus; (ii) establishes a potential mechanism of simultaneous tongue protrusion and larynx retraction by applying a two-dimensional model based on graphic reconstructions in single video frames of unrestrained animals; and (iii) advances a hypothesis of evaporative cooling by tongue protrusion in the males of a subspecies of red deer constrained to perform all of the exhausting rutting activities, including acoustic display, in a hot and arid season.

Descended and mobile larynx, vocal tract elongation and rutting roars in male goitred gazelles (Gazella subgutturosa Güldenstaedt, 1780).

Similar to male humans, Homo sapiens, the males of a few polygynous ruminants - red deer Cervus elaphus, fallow deer Dama dama and Mongolian gazelle Procapra gutturosa- have a more or less enlarged, low-resting larynx and are capable of additional dynamic vocal tract elongation by larynx retraction during their rutting calls. The vocal correlates of a large larynx and an elongated vocal tract, a low fundamental frequency and low vocal tract resonance frequencies, deter rival males and attract receptive females. The males of the polygynous goitred gazelle, Gazella subgutturosa, provide another, independently evolved, example of an enlarged and low-resting larynx of high mobility. Relevant aspects of the rutting behaviour of territorial wild male goitred gazelles are described. Video and audio recordings served to study the acoustic effects of the enlarged larynx and vocal tract elongation on male rutting calls. Three call types were discriminated: roars, growls and grunts. In addition, the adult male vocal anatomy during the emission of rutting calls is described and functionally discussed using a 2D-model of larynx retraction. The combined morphological, behavioural and acoustic data are discussed in relation to the hypothesis of sexual selection for male-specific deep voices, resulting in convergent features of vocal anatomy in a few polygynous ruminants and in human males.

Developmental changes of nasal and oral calls in the goitred gazelle Gazella subgutturosa, a nonhuman mammal with a sexually dimorphic and descended larynx.

In goitred gazelles (Gazella subgutturosa), sexual dimorphism of larynx size and position is reminiscent of the case in humans, suggesting shared features of vocal ontogenesis in both species. This study investigates the ontogeny of nasal and oral calls in 23 (10 male and 13 female) individually identified goitred gazelles from shortly after birth up to adolescence. The fundamental frequency (f0) and formants were measured as the acoustic correlates of the developing sexual dimorphism. Settings for LPC analysis of formants were based on anatomical dissections of 5 specimens. Along ontogenesis, compared to females, male f0 was consistently lower both in oral and nasal calls and male formants were lower in oral calls, whereas the first two formants of nasal calls did not differ between sexes. In goitred gazelles, significant sex differences in f0 and formants appeared as early as the second week of life, while in humans they emerge only before puberty. This result suggests different pathways of vocal ontogenesis in the goitred gazelles and in humans.

Hiss and snort call types of wild-living giraffes Giraffa camelopardalis: acoustic structure and context.

OBJECTIVES: Vocalization as part of vigilance behaviour is widespread across animal taxa, including ruminants. Calls of wild-living giraffes have never been recorded and spectrographically investigated. This study reports the acoustic structure of vigilance-related hiss and snort calls of wild-living giraffes Giraffa camelopardalis. RESULTS: The hiss and snort calls were emitted during five recording sessions produced by nine individual giraffes (8 adults and 1 subadult) in their natural environment in Namibia (3 individuals) and Kenya (6 individuals). These calls attended vigilance behaviour toward humans in hides or in vehicles and cheetahs as natural predators of giraffe young. This study provides spectrographic analyses of 22 hiss and 20 snort calls. The giraffe hisses were broadband vocalizations of an average duration of 0.72 s (from 0.24 to 1.04 s) and a peak frequency of 0.69 kHz. The giraffe snorts were broadband pulsed calls of an average duration of 0.28 s (from 0.13 to 0.55 s), a peak frequency at 0.20 kHz and comprised a prominent low-frequency pulsation of 23.7 pulses/s. The acoustic structure of giraffe hisses is reminiscent of vigilance-related hisses of musk deer Moschus moschiferus. Giraffe snorts differ from snorts of other ruminants by their prominent pulsed pattern.

Vocal changes accompanying the descent of the larynx during ontogeny from neonates to adults in male and female goitred gazelles (Gazella subgutturosa).

The pronouncedly enlarged and descended larynx in male goitred gazelles (Gazella subgutturosa), Mongolian gazelles (Procapra gutturosa) and fallow deer (Dama dama) represents an interesting parallel to the 'Adam's apple' of human males. Goitred gazelles, as humans, are not born with a descended larynx. Therefore the sexual dimorphism of larynx size and position develops during ontogeny. In this study, the vocal ontogeny of male and female goitred gazelles was investigated across five age classes from neonates to adults. The acoustic variables of nasal contact calls were measured in 53 (24 male, 29 female) individuals, body mass and neck dimensions in 63 (31 male, 32 female) live individuals and nasal vocal tract and vocal fold lengths in 26 (16 male, 10 female) anatomical specimens. Call fundamental frequency (f0), the acoustic correlate of the ontogenetically enlarging larynx, decreased significantly in either sex. Call formants (second, third and forth), the acoustic correlates of the ontogenetically elongating vocal tract, did not differ significantly between sexes up to early adulthood, but clearly diverged in adults. Significant differences between sexes in neck circumference at the level of the larynx emerged already at 2-3 months of age, whereas body mass, neck circumference at the neck-body transition and the degree of larynx descent significantly differed in adults only. We discuss that, in contrast to humans, the accelerated enlargement of the larynx in male goitred gazelles starts early in ontogeny. A moderate descent of the larynx develops equally in both sexes before early adulthood, whereas the additional prominent descent of the larynx in males is shifted to late ontogeny. This might avoid selective disadvantages of this sexually dimorphic trait on males during their period of growth. As has been previously proved for humans, the emergence of the strong male-specific descent of the larynx in goitred gazelles may go along with the increasing social status and the males' increasing chances of siring offspring. Similar to the Adam's apple of human males, this may indicate the important role of the enlarged and descended larynx for signaling male status via masculine voice.

Potential Sources of High Frequency and Biphonic Vocalization in the Dhole (Cuon alpinus).

Biphonation, i.e. two independent fundamental frequencies in a call spectrum, is a prominent feature of vocal activity in dog-like canids. Dog-like canids can produce a low (f0) and a high (g0) fundamental frequency simultaneously. In contrast, fox-like canids are only capable of producing the low fundamental frequency (f0). Using a comparative anatomical approach for revealing macroscopic structures potentially responsible for canid biphonation, we investigated the vocal anatomy for 4 (1 male, 3 female) captive dholes (Cuon alpinus) and for 2 (1 male, 1 female) wild red fox (Vulpes vulpes). In addition, we analyzed the acoustic structure of vocalizations in the same dholes that served postmortem as specimens for the anatomical investigation. All study dholes produced both high-frequency and biphonic calls. The anatomical reconstructions revealed that the vocal morphologies of the dhole are very similar to those of the red fox. These results suggest that the high-frequency and biphonic calls in dog-like canids can be produced without specific anatomical adaptations of the sound-producing structures. We discuss possible production modes for the high-frequency and biphonic calls involving laryngeal and nasal structures.

The postnatal ontogeny of the sexually dimorphic vocal apparatus in goitred gazelles (Gazella subgutturosa).

This study quantitatively documents the progressive development of sexual dimorphism of the vocal organs along the ontogeny of the goitred gazelle (Gazella subgutturosa). The major, male-specific secondary sexual features, of vocal anatomy in goitred gazelle are an enlarged larynx and a marked laryngeal descent. These features appear to have evolved by sexual selection and may serve as a model for similar events in male humans. Sexual dimorphism of larynx size and larynx position in adult goitred gazelles is more pronounced than in humans, whereas the vocal anatomy of neonate goitred gazelles does not differ between sexes. This study examines the vocal anatomy of 19 (11 male, 8 female) goitred gazelle specimens across three age-classes, that is, neonates, subadults and mature adults. The postnatal ontogenetic development of the vocal organs up to their respective end states takes considerably longer in males than in females. Both sexes share the same features of vocal morphology but differences emerge in the course of ontogeny, ultimately resulting in the pronounced sexual dimorphism of the vocal apparatus in adults. The main differences comprise larynx size, vocal fold length, vocal tract length, and mobility of the larynx. The resilience of the thyrohyoid ligament and the pharynx, including the soft palate, and the length changes during contraction and relaxation of the extrinsic laryngeal muscles play a decisive role in the mobility of the larynx in both sexes but to substantially different degrees in adult females and males. Goitred gazelles are born with an undescended larynx and, therefore, larynx descent has to develop in the course of ontogeny. This might result from a trade-off between natural selection and sexual selection requiring a temporal separation of different laryngeal functions at birth and shortly after from those later in life. J. Morphol. 277:826-844, 2016. © 2016 Wiley Periodicals, Inc.

Vocal development during postnatal growth and ear morphology in a shrew that generates seismic vibrations, Diplomesodon pulchellum.

The ability of adult and subadult piebald shrews (Diplomesodon pulchellum) to produce 160Hz seismic waves is potentially reflected in their vocal ontogeny and ear morphology. In this study, the ontogeny of call variables and body traits was examined in 11 litters of piebald shrews, in two-day intervals from birth to 22 days (subadult), and ear structure was investigated in two specimens using micro-computed tomography (micro-CT). Across ages, the call fundamental frequency (f0) was stable in squeaks and clicks and increased steadily in screeches, representing an unusual, non-descending ontogenetic pathway of f0. The rate of the deep sinusoidal modulation (pulse rate) of screeches increased from 75Hz at 3-4 days to 138Hz at 21-22 days, probably relating to ontogenetic changes in contraction rates of the same muscles which are responsible for generating seismic vibrations. The ear reconstructions revealed that the morphologies of the middle and inner ears of the piebald shrew are very similar to those of the common shrew (Sorex araneus) and the lesser white-toothed shrew (Crocidura suaveolens), which are not known to produce seismic signals. These results suggest that piebald shrews use a mechanism other than hearing for perceiving seismic vibrations.

The anatomy of vocal divergence in North American Elk and European red deer.

Loud and frequent vocalizations play an important role in courtship behavior in Cervus species. European red deer (Cervus elaphus) produce low-pitched calls, whereas North American elk (Cervus canadensis) produce high-pitched calls, which is remarkable for one of the biggest land mammals. Both species engage their vocal organs in elaborate maneuvers but the precise mechanism is unknown. Vocal organs were compared by macroscopic and microscopic dissection. The larynx is sexually dimorphic in red deer but not in elk. The laryngeal lumen is more constricted in elk, and narrows further during ontogeny. Several elements of the hyoid skeleton and two of four vocal tract segments are longer in red deer than in elk allowing greater vocal tract expansion and elongation. We conclude that elk submit the larynx and vocal tract to much higher tension than red deer, whereby, enormously stressed vocal folds of reduced effective length create a high resistance glottal source. The narrow, high impedance laryngeal vestibulum matches glottal and vocal tract impedance allowing maximum power transfer. In red deer longer and relaxed vocal folds create a less resistant glottal source and a wider vestibulum matches the low glottal impedance to the vocal tract, thereby also ensuring maximum power transfer.

Koalas use a novel vocal organ to produce unusually low-pitched mating calls.

During the breeding season, male koalas produce 'bellow' vocalisations that are characterised by a continuous series of inhalation and exhalation sections, and an extremely low fundamental frequency (the main acoustic correlate of perceived pitch) [1]. Remarkably, the fundamental frequency (F0) of bellow inhalation sections averages 27.1 Hz (range: 9.8-61.5 Hz [1]), which is 20 times lower than would be expected for an animal weighing 8 kg [2] and more typical of an animal the size of an elephant (Supplemental figure S1A). Here, we demonstrate that koalas use a novel vocal organ to produce their unusually low-pitched mating calls.

How low can you go? Physical production mechanism of elephant infrasonic vocalizations.

Elephants can communicate using sounds below the range of human hearing ("infrasounds" below 20 hertz). It is commonly speculated that these vocalizations are produced in the larynx, either by neurally controlled muscle twitching (as in cat purring) or by flow-induced self-sustained vibrations of the vocal folds (as in human speech and song). We used direct high-speed video observations of an excised elephant larynx to demonstrate flow-induced self-sustained vocal fold vibration in the absence of any neural signals, thus excluding the need for any "purring" mechanism. The observed physical principles of voice production apply to a wide variety of mammals, extending across a remarkably large range of fundamental frequencies and body sizes, spanning more than five orders of magnitude.

The ontogeny of acoustic individuality in the nasal calls of captive goitred gazelles, Gazella subgutturosa.

Individualistic voices are important for establishing personalized relationships among individuals. In young animals, individual vocal identity is affected by permanent changes of the acoustics due to the growth of their vocal apparatus. Different acoustic variables change uncoordinatedly, so vocal individuality should be repeatedly upgraded along development. We compared classifying accuracy of individuals and sexes by nasal calls in fast-growing goitred gazelles Gazella subgutturosa at two ontogenetic stages, juvenile (3-6 weeks of age) and adolescent (23-26 weeks of age). Juvenile "spring" nasal calls and adolescent "fall" nasal calls were examined in the same 35 calves (18 males, 17 females), wild-born in May and then hand-raised. Discriminate function analysis based on four formants, fundamental frequency, duration and three power quartiles, revealed an equally high potential of spring and fall calls to encode sex. The individuality was very high in both ages but significantly higher in fall calls. Classifying calls to individuals was based on the same three acoustic variables (fundamental frequency and third and fourth formants) in both ages, although their actual values changed uncoordinatedly from spring to fall in most subjects. Our results suggest updating acoustic individuality in nasal calls of adolescent goitred gazelles accordingly to the newly emerged acoustic variation.