Alterations in the release calls of six European anura (amphibia) after partial or total extirpation of the vocal cords

In the interior of the larynx of Ranidae there are two sturdy vocal cords. The Bufonidae have more delicate vocal cords, and in addition paired cushions of tissue anterior to the cords and paired folds posterior to the cords.In the three ranids Rana esculenta, Rana ridibunda and Rana temporaria, partial or total extirpation of the vocal cords results in loss of voice or atypical release calls. In such remnants of calls as are retained, the frequency composition is little affected, whereas the intensity is always greatly reduced. The most severe impairment is evident in the formation of sound pulses and in the rhythmicity of the pulse sequence.In the three bufonids Bufo bufo, Bufo calamita and Bufo viridis loss of voice is a less common result of the various operations than in the ranids. The most marked deterioration follows removal of all or part of the vocal cords. The tissue cushions and the posterior folds participate, along with the vocal cords, in production of the release calls. Post-operative alterations in the release calls are therefore quite variable.     

The vocal sac increases call rate in the Tungara frog Physalaemus pustulosus

In most anurans, the production of advertisement calls is accompanied by the inflation of a vocal sac. Current functions of the vocal sac, however, are not fully understood, although several hypotheses have been proposed. One hypothesis suggests that the vocal sac decreases the intercall interval (i.e., increases call rate) by reinflating the lungs more rapidly than is possible with the buccal pump. We investigate this hypothesis by analyzing audio and video recordings of calling tungara frogs. We compare the first two call bouts emitted by an originally uninflated male. The first call bout requires lung inflation via buccal pumping, but in the second, the male is already inflated because of capture of air and reinflation of the lungs by the vocal sac. Lung inflation to typical field levels requires 26-51 buccal pumps, which takes at least 4.4 s. This estimate is more than 2.5 times the typical intercall interval with lung reinflation via a vocal sac (ca. 1.7 s). Evidence from phonotaxis tests demonstrates that these differences in intercall intervals are salient to females and that female Physalaemus pustulosus prefer the shorter intercall interval/higher call rate. Acoustic analyses demonstrate that the first call of bout 1, which requires buccal pumping, is usually shorter, of lower amplitude, and spans a smaller frequency range than the first call of bout 2, which does not require buccal pumping. Because females prefer longer, more intense calls, these results suggest that the vocal sac not only increases call rate but also allows males to produce more calls of increased attractiveness to females.     

Muscular control of the vocal tract during release signaling in the toad Bufo valliceps

The release vibration and release call of Bufo valliceps have been studied by electromyography of the muscles involved, coupled with pressure and sound recording. The sequences are powered by contraction of the muscles of the body envelope and with the energy transmitted via the compressed pulmonary contents. Each pulse of a call starts as the laryngeal muscles relax and pulmonary pressure forces the arytenoid cartilages apart. Sound emission ceases when the laryngeal dilators pull the arytenoids out of the airstream. Reverse flow of air from buccal cavity to lungs may occur within prolonged release sequences. Inflation of the vocal sac results in marked increase in amplitude of the radiated sound without equivalent increase in amplitude of the myograns. The call is intimately associated with the pulsepumping method of breathing used by frogs.     

Sexually dimorphic laryngeal morphology in Rana pipiens

The sexually dimorphic vocal characteristics of Rana pipiens release calls suggest that there may be differences in the anatomical components of the larynx. The volumes of the arytenoid cartilage, surrounding muscle masses, vocal cords, supporting bronchial process, and the release-call amplitudes of six males and five females were measured in same-sized animals and sexual differences assessed. No qualitative differences in laryngeal morphology were observed, but all features measured except vocal cords were significantly larger in males. The implications of an increased laryngeal size are discussed in relation to differences previously observed in the vocalizations of this and other species and in relation to prior suggestions regarding the developmental basis of anuran sexual dimorphisms.     

Vocal morphology of the Physalaemus pustulosus species group (Leptodactylidae): morphological response to sexual selection for complex calls

Most male frogs in the genus Physalaemus produce a whine-like advertisement call. Male P. pustulosus , however, add chucks to the call. This enhances the attractiveness of the call to females, and has evolved under the influence of sexual selection despite the increased predation risk from the frog-eating bat ( Trachops cirrhosus ). This complex call is unusual, if not unique, among anurans because the two call components overlap in time. Here we investigate the morphological changes responsible for the production of complex calls.
The Physalaemus purtulosus species group consists of four species. Physalaemus pustulosus and P. petersi are sister species, and recently it has been shown that P. petersi produces chucks. Physalaemus coloradorum and P. purtulatus are sister species and neither is known to produce chucks. Two laryngeal characters vary within the species group. Physalaemus pustulosus has a large fibrous mass (FMI), whose vibration is responsible for production of the chuck. This mass is much smaller in the other three species. In P. pustulosus and P. petersi the FMI is anchored dorsally, deep within the bronchial process, the attachment is more extensive in P. pustulosus. Neither P. pustulatus nor P. coloradorum have such a dorsal attachment associated with their FMI. This character is responsible for allowing the FMI to vibrate independently of the vocal cords, that is, for the production of the complex call. Thus the morphological changes responsible for the evolution of this unusual behavioural innovation, the complex call, are gradual, and almost trival, in nature. This study also shows that the primitive condition of the larynx of the P. pustulosus and P. petersi ancestor, was predisposed to the production of complex calls.
We also document ontogenetic and sexually dimorphic patterns in larynx structure.     

Vocal production in nonhuman primates: Acoustics,physiology, and functional constraints on “honest” advertisement

The physiological mechanisms and acoustic principles underlying sound production in primates are important for analyzing and synthesizing primate vocalizations, for determining the range of calls that are physically producible, and for understanding primate communication in the broader comparative context of what is known about communication in other vertebrates. In this paper we discuss what is known about vocal production in nonhuman primates, relying heavily on models from speech and musical acoustics. We first describe the role of the lungs and larynx in generating the sound source, and then discuss the effects of the supralaryngeal vocal tract in modifying this source. We conclude that more research is needed to resolve several important questions about the acoustics of primate calls, including the nature of the vocal tract's contribution to call production. Nonetheless, enough is known to explore the implications of call acoustics for the evolution of primate communication. In particular, we discuss how anatomy and physiology may provide constraints resulting in “honest” acoustic indicators of body size. © 1995 Wiley-Liss, Inc.     

Food-associated calls in rhesus macaques (Macaca mulatta): I. Socioecological factors

Upon discovering food, free-living rhesus macaques (Macaca mulatta)on the island of Cayo Santiago, Puerto Rico, produce a complexof vocal signals consisting of five acoustically distinguishablecalls. This report examines the socioecological factors elicitingcall production and the information protentially conveyed toothers. The primary contexts for three vocalizations ("warbles," "harmonicarches, " and "chirps") are encounters with rareand highly preferred foods (e.g., coconut). Two other vocalizations("coos" and "grunts") are produced both in food (primarily provisionedchow) and in nonfood contexts, such as during mother-infantseparation and grooming interactions. Grunts given upon encounteringfood are acoustically distinct from those given in nonfood contexts.In contrast, coos associated with food are statistically indistinguishablefrom coos given in other contexts. When conspecitics hear thesefood-associated calls, they typically approach the caller. Coosare less likely to lead to approach than other food-associatedcalls, Results from all-day follows on adult males and adultfemales reveal that changes in hunger level influence call ratebut not call type; the different call types are produced throughoutthe day. We infer that the structure of food-associated callsprovides information about the quality of the food discovered,whereas call rate conveys information about the relative hungerlevel of the caller. In this population, adult males give fewerfood-associated calls than adult females. In addition, femaleswithin large matrilines call more than females within smallermatrilines, and males who are resident in a group are more vocalthan peripheral males.     

Efficiency and Significance of Multiple Vocal Signals in Sibling Competition

Animals can compete for resources by displaying various acoustic signals that may differentially affect the outcome of competition. We propose the hypothesis that the most efficient signal to deter opponents should be the one that most honestly reveals motivation to compete. We tested this hypothesis in the barn owl (Tyto alba) in which nestlings produce more calls of longer duration than siblings to compete for priority access to the indivisible prey item their parents will deliver next. Because nestlings increase call rate to a larger extent than call duration when they become hungrier, call rate would signal more accurately hunger level. This leads us to propose three predictions. First, a high number of calls should be more efficient in deterring siblings to compete than long calls. Second, the rate at which an individual calls should be more sensitive to variation in the intensity of the sibling vocal competition than the duration of its calls. Third, call rate should influence competitors’ vocalization for a longer period of time than call duration. To test these three predictions we performed playback experiments by broadcasting to singleton nestlings calls of varying durations and at different rates. According to the first prediction, singleton nestlings became less vocal to a larger extent when we broadcasted more calls compared to longer calls. In line with the second prediction, nestlings reduced vocalization rate to a larger extent than call duration when we broadcasted more or longer calls. Finally, call rate had a longer influence on opponent’s vocal behavior than call duration. Young animals thus actively and differentially use multiple signaling components to compete with their siblings over parental resources.     

Experimental study of alarm calls of the oriental tit (Parus minor) toward different predators and reactions they induce in nestlings

Anti-predatory strategies of birds are diverse and may include predator-specific alarm calls. For example, oriental tit (Parus minor) parents can distinguish snakes from other predators and produce snake-specific referential vocalizations ("jar" call) when a snake poses a threat to their nest. The “jar” call has a very specific function to induce fledging of nestlings close to fledging age. This reaction ensures nestlings' survival in natural encounters with snakes that are capable of entering nest cavities and kill entire broods. Sciurid rodents, like chipmunks, may pose a similar threat to cavity-nesting birds. We explored the hypothesis that parents use the fledging-inducing alarm vocalizations in this situation, because chipmunks, like snakes, can kill the brood upon entering the nest cavity. We compared alarm calls of parents toward two predators (chipmunk and snake) who pose a similar threat to the nestlings in a nest cavity, and toward an avian predator (Eurasian jay) who cannot enter nest cavities and poses no threat to the nestlings in a nest. Our results show that the vocal responses of oriental tits were different among the three predators. This suggests that the acoustic properties of vocal responses to predators are different between predators of a similar hunting strategy (nest-cavity entering). The playback of recorded vocal responses of parents to chipmunks did not trigger the fledging of old nestlings, whereas the vocalizations toward a snake did, as shown by earlier studies. Our study suggests that the vocal response of parents does not carry information about the ability of predators to enter the nest cavity and confirms the special status of alarm calls triggered by snakes.     

Vocal Tract Morphology Determines Species-Specific Features in Vocal Signals of Lemurs (Eulemur)

The source-filter theory describes vocal production as a two-stage process involving the generation of a sound source, with its own spectral structure, which is then filtered by the resonant properties of the vocal tract. This theory has been successfully applied to the study of animal vocal signals since the 1990s. As an extension, models reproducing vocal tract resonance can be used to reproduce formant patterns and to understand the role of vocal tract filtering in nonhuman vocalizations. We studied three congeneric lemur species —Eulemur fulvus, E. macaco, E. rubriventer— using morphological measurements to build computational models of the vocal tract to estimate formants, and acoustic analysis to measure formants from natural calls. We focused on call types emitted through the nose, without apparent articulation. On the basis of anatomical measurements, we modeled the vocal tract of each species as a series of concatenated tubes, with a cross-sectional area that changed along the tract to approximate the morphology of the larynx, the nasopharyngeal cavity, the nasal chambers, and the nostrils. For each species, we calculated the resonance frequencies in 2500 randomly generated vocal tracts, in which we simulated intraspecific length and size variation. Formant location and spacing showed significant species-specific differences determined by the length of the vocal tract. We then measured formants of a set of nasal vocalizations (“grunts”) recorded from captive lemurs of the same species. We found species-specific differences in the natural calls. This is the first evidence that morphology of the vocal tract is relevant in generating filter-related acoustic cues that potentially provide receivers with information about the species of the emitter.     

Structure of the larynx of the tokay gecko (Gekko gecko), with particular reference to the vocal cords and glottal lips

The ability to vocalize is well-known in gekkonid lizards but relatively little attention has been paid to the structure of the vocal apparatus. In this study we briefly review the structure of the larynx and associated musculature of the tokay gecko as a baseline for a comparative survey of the family. The cricoid and arytenoid cartilages form the skeleton of the larynx and are controlled by constrictor and dilator muscles. The gross morphology of the vocal cords and glottal lips is then described, the structure being elucidated by way of dissection, histology, and scanning electron microscopy. The vocal cords run between the arytenoid and cricoid cartilages, are highly elastic, and bear a highly folded mucosa. The lips of the glottis have a structure reminiscent of erectile tissue. The respiratory mucosa of the larynx varies according to position and may be related to the tonal aspects of sound production. The structure of the larynx is compared with that of other vertebrates, and the relationship between morphology and phonation is considered.     

Vocal repertoire ontogeny of the captive Asian house shrew Suncus murinus suggests that the male courtship call develops from the caravanning call of the young

Soricids produce a considerable variety of vocalizations. However, these calls have been studied insufficiently with the exception of echolocation calls. In this study, 1,645 calls from 18 juvenile, ten sub-adult and 36 adult Asian house shrews (Suncus murinus) were acoustically and statistically analyzed to describe this species’ vocal repertoire and its ontogeny. The vocal repertoire of S. murinus includes 17 call types, seven tonal (whistle, chirp, twitter, whimper, squeak, scream and short scream) and ten non-tonal (churr, shriek, babble, click, boom, snort, screech, short screech, sniff and low click), of which ten call types (whimper, squeak, scream, short scream, churr, babble, snort, short screech, sniff and low click) were newly described by this study. This relatively extensive vocal repertoire, including one call type emitted during collective resting, indicates that this species possibly possesses a higher degree of sociality and cohesiveness than previously expected. High structural similarities were observed between calls produced by juveniles and sub-adults during caravanning and those produced by adult males during courtship. Therefore, the results of this study support a previously suggested hypothesis that in shrews, adult courtship calls are derived from calls emitted by the young. The results of this study also showed that the largest changes in the ontogeny of the vocal repertoire occurred at approximately 10 days old and was in close connection to the eyes opening. The results are discussed with available information on the vocal repertoires of other soricids.     

No need to shout: Effect of signal loudness on sibling communication in barn owls Tyto alba

In animal communication, signal loudness is often ignored and seldom measured. We used a playback experiment to examine the role of vocal loudness (i.e., sound pressure level) in sibling to sibling communication of nestling barn owls Tyto alba. In this species, siblings vocally negotiate among each other for priority access to parental food resources. Call rate and call duration play key roles in this vocal communication system, with the most vocal nestlings deterring their siblings from competing for access to the food item next delivered by parents. Here, we broadcast calls at different loudness levels and call rate to live nestlings. The loudness of playback calls did not affect owlets' investment in call rate, call duration or call loudness. The rate at which playback calls were broadcast affected owlets' call rate but did not influence their response in terms of loudness. This suggests that selection for producing loud signals may be weak in this species, as loud calls may attract predators. Moreover, given that owlets do not overlap their calls and that they communicate to nearby siblings in the silence of the night, loud signals may not be necessary to convey reliable information about food need.     

Release vocalizations in neotropical toads (Bufo): ecological constraints and phylogenetic implications

The release vocalizations of four nominal neotropical toad taxa ( Bufo achalensis , Bufo limensis , Bufo spinulosus , Bufo arenarum ) which compose three sympatric species pairs, were quantitatively analysed and homologous call types statistically compared. The first three taxa are closely related members of the mainly Andean Bufo spinulosus species group in which advertisement calls are absent. The specific vocal repertoire consisted of a uniform, unpulsed release call and one (in B. arenarum, two) pulsed release trill(s) which were given either singly or in series of up to three single calls. The uniform call was similar in structure and dominant (= basic) frequency in all taxa and probably represents an unspecific acoustic signal which is used to avoid heterospecific amplexus between sympatric toads. The release trills significantly differed in most call features among the taxa and, using discriminant analyses, even single calls were correctly assigned to sympatric pairs of species. In contrast, calls of allopatric pairs of species were confounded at rates of up to 36% indicating that selection towards species-specific signals increases when different species live in sympatry. The release trills of two allopatric populations (Perú, Argentina) assigned to B. spinulosus were similar in structure, but nevertheless features of pulse train permitted an unequivocal distinction, suggesting a long-lasting independent evolution. The taxonomic significance of this finding remains to be evaluated in further investigations. The complex vocal repertoire also offered the opportunity to assess phylogenetic relationships among the taxa. The extra-Andean B. achalensis seems to be closer related to the Andean B. spinulosus than this species is to the widely sympatric B. limensis – a species with several primitive character states indicating an early separation from the ancestral stock.     

Vocal repertoire of cooperatively breeding Smooth‐billed Anis

Calls are functionally diverse signals that mediate behavior in a wide variety of contexts in both passerines and non‐passerines. However, the call‐based acoustic communication systems of non‐passerines have received less attention from investigators than those of passerines. We examined the vocal repertoire of Smooth‐billed Anis (Crotophaga ani), cooperatively breeding cuckoos that live in groups with multiple breeding pairs. We recorded calls from 22 groups over two breeding seasons at the Cabo Rojo National Wildlife Refuge in Puerto Rico. We identified 11 call types and one group vocalization, and used an automated sound measurement program to quantify their acoustic features. Discriminant function analysis (DFA) correctly classified 74.2% of calls based on these features. The vocal repertoire of Smooth‐billed Anis is larger than that reported for the three other species in the subfamily Crotophaginae. Smooth‐billed Anis have at least two alarm calls, two nest‐specific calls, and one nest defense call. We also identified one possible signal of aggressive intent, one possible appeasement signal, and two calls that may communicate identity. The relatively large vocal repertoire of Smooth‐billed Anis and association of distinct call types with different functions and contexts supports the main prediction of the social complexity hypothesis, i.e., species with more complex social systems will have more complex communication systems.     

Vasotocin maintains multiple call types in the gray treefrog, Hyla versicolor.

The neuropeptide arginine vasotocin (AVT) influences vocalizations in some anuran amphibians but it is unknown whether AVT alters all vocal behaviors of a species similarly. We first characterized the vocal repertoire of male gray treefrogs (Hyla versicolor). Three different call types were distinguished by unique sets of temporal and spectral features. Second, we examined the effects of AVT on each call type by injecting frogs with either AVT (100 microg; intraperitoneal) or saline and recording subsequent behavior. In the field, AVT maintained advertisement calling, whereas calling ceased in saline-injected animals. Advertisement call rate in AVT-injected males fell significantly and dominant frequency of the call was significantly higher. In the laboratory, AVT induced advertisement calling in males that were not initially vocalizing and dominant frequency was also significantly higher in these males. AVT maintained aggressive calling similarly but the characteristics of aggressive calls were not altered by AVT. There were no significant differences in release call behavior between AVT- and saline-injected groups; however, release call duration decreased significantly in AVT-injected animals, compared with preinjection values for the same animals. The effects of AVT on vocal behavior in this species are therefore not the same for each call type. AVT may act at more general motivational levels in the central nervous system and other neural or endocrine factors may control choice of call type and direct motor output.     

四川梅花鹿产仔换茸期声行为的研究

2009年4～9月在四川省若尔盖县铁布自然保护区,对四川梅花鹿Cervusnippon sichuanicus产仔换茸期的声行为进行了较系统的研究。四川梅花鹿产仔换茸期的声行为可以分为报警声、惊叫声、吼叫声、母子召唤声等20种鸣声。通过声谱分析获得了各种鸣声的语谱图及其频谱特征,并通过野外观察初步确定了这些声行为的生物学意义。在遇到危险或潜在危险时,成年雄鹿往往第一个发出报警鸣声,而雌鹿的报警声大约是雄鹿的两倍多。成年个体的主动防御鸣声频次要多于亚成体,而幼体未见有该类鸣声。对梅花鹿不同亚种和不同季节的几种声行为进行了比较。     

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. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.