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Howling

Howling is a prolonged, melodious vocalization produced by various mammals, primarily canids such as (Canis lupus) and domestic dogs (Canis familiaris), as well as howler monkeys (genus Alouatta), serving as a primary form of long-distance communication to maintain social bonds, locate separated group members, and defend territories. In canids, howling functions as a contact call that facilitates pack reassembly after separations, often during hunts or dispersals, and is influenced by social relationships rather than physiological stress levels like . Studies have identified distinct "dialects" in howls across species and , with acoustic variations in and that encode individual identity, group , and even genetic relatedness. These vocalizations can propagate over several kilometers, enabling coordination in low-visibility environments, and are typically contagious within groups, where one individual's prompts others to join. Among howler monkeys, —often characterized as powerful roars—represents one of the loudest vocalizations, audible up to 5 kilometers away, and is predominantly produced by males to advertise presence, deter intruders, and attract mates. Evolutionary trade-offs influence these calls; for instance, larger hyoid bones in males enhance roar volume but correlate with reduced testes size due to energy allocation constraints. in these peaks at dawn and dusk, aligning with territorial defense and group cohesion in dense forest habitats. Beyond these prominent examples, howling occurs in other species like coyotes and certain foxes, though less prominently, and shares acoustic similarities that suggest for signaling in open or obstructed environments. Research continues to explore the neural and genetic bases of howling, revealing its role in and across taxa.

Overview and Definition

Definition of Howling

is a sustained, resonant produced by the vibration of the vocal folds in the , generating a prolonged, often undulating that facilitates long-distance communication among . This acoustic involves controlled and modulation of , distinguishing it as a form of adapted for propagation over extended ranges in various environments. Unlike barking, which consists of short, abrupt bursts typically used for immediate alerts or territorial warnings, howling features a more continuous and melodic quality with variable pitch transitions. In contrast, roaring—common in large felids—relies on lower-frequency, explosive bursts enabled by specialized, square-shaped vocal folds that produce intense, low-pitched rumbles for or displays. howls exemplify the melodic variability of , often rising and falling in pitch to convey emotional or . The term "howling" derives from the Middle English "houlen," an imitative word from the early meaning to emit a loud, prolonged cry, akin to similar forms in other and tracing back to an Old English root "*hūlian" referring to extended vocal cries. Howling is prevalent among certain mammals, particularly carnivores such as canids, and including howler monkeys, where it serves adaptive roles in social and ecological contexts, though it is not observed universally across mammalian species.

Acoustic Characteristics

Howling sounds are characterized by their low fundamental frequencies, which facilitate long-distance transmission. In canines such as wolves, the fundamental frequency typically ranges from 300 to 1000 Hz, with harmonics extending up to approximately 2000 Hz, concentrating acoustic energy in the lower spectrum for efficient propagation. In howler monkeys, vocalizations feature emphasized frequency bands starting as low as 200-700 Hz, with peaks often reaching 800-1200 Hz due to their enlarged hyoid apparatus, which lowers the formant dispersion. The duration of individual howls generally spans 5 to 20 seconds, though chorus events involving multiple animals can extend much longer, averaging around 66 seconds in packs. These vocalizations often exhibit a song-like structure, with rising and falling pitches that include frequency modulations and discontinuities, creating undulating patterns visible in acoustic analyses. Low-frequency components of howls experience minimal , allowing up to 10 km in open such as , where calm air enhances transmission. Environmental factors like , which can scatter higher frequencies, and , such as forests that increase , significantly reduce effective range, sometimes limiting detection to 6 km or less in denser habitats. Spectrograms are commonly employed to measure howling acoustics, revealing formants as dark bands corresponding to resonant frequencies and tracking pitch variations through fundamental frequency contours. For instance, studies on wolf howls report average source amplitudes of 90-110 , with energy concentrated in the initial harmonics, enabling differentiation of individual and group contributions.

Biological and Physiological Aspects

Mechanisms of Sound Production

Howling is produced through the coordinated of the respiratory and laryngeal systems, where air expelled from the lungs passes through the to vibrate the vocal folds, generating waves. The , located at the top of the trachea, houses the vocal folds—pairs of muscular membranes that can be tensed, relaxed, or approximated by surrounding cartilages and muscles to control pitch and volume. In many howling species, such as howler monkeys (Alouatta spp.), the vocal folds are notably elongated relative to body size, measuring up to 4.08 cm in length, which enables the production of low-frequency, resonant calls. Throat muscles, including the extrinsic laryngeal muscles attached to the , modulate the tension and position of the vocal folds, allowing precise adjustments to the during sustained howls. A key anatomical feature enhancing howl production is the , which suspends the and contributes to resonance. In howler monkeys, the hyoid bone is greatly enlarged, forming a bulla-like chamber that acts as a , amplifying the sound by reflecting and intensifying vibrations from the vocal folds. This , enclosed by deep mandibles, can increase call intensity significantly, enabling howls to travel over 3 km in forested environments. Neural control of these processes originates in the , particularly the , which innervates laryngeal muscles via the vagus and glossopharyngeal nerves, while higher modulation from the integrates emotional and motivational inputs to initiate and sustain vocalization. The in the serves as a central , coordinating brainstem outputs for rhythmic air expulsion and fold vibration. Variations in sound production mechanisms exist across taxa, reflecting adaptations to ecological demands. In canines like wolves (Canis lupus), the epiglottis plays a crucial role in resonance by altering airflow dynamics and contributing to the harmonic structure of howls; adjustments in its angle can enhance acoustic output in excised larynges. Primates such as howler monkeys and siamangs (Symphalangus syndactylus) utilize laryngeal air sacs or enlarged hyoid structures—functionally akin to throat pouching—for additional resonance and prolonged phonation without excessive lung strain. These sacs store exhaled air, allowing rapid call sequences while minimizing hyperventilation risks. Howling demands substantial energy, involving forceful diaphragmatic contractions and sustained subglottal pressure, which can represent a notable portion of daily metabolic expenditure in smaller species with frequent vocal bouts. Pathological conditions can disrupt these mechanisms, notably in dogs, where degeneration of the impairs abductor muscle function, leading to incomplete opening of the vocal folds and reduced airflow for . This results in hoarse, weakened, or absent vocalizations, including the inability to produce normal howls, alongside respiratory distress. Surgical interventions, such as arytenoid lateralization, aim to widen the rima glottidis and restore partial phonatory capacity.

Functions and Evolutionary Role

Howling serves several primary adaptive functions in social mammals, particularly in species inhabiting dense or forested environments where visual cues are limited. These include territory defense, pack or group coordination, mate attraction, and distress signaling. In wolves (Canis lupus), howling functions to advertise pack presence and boundaries, promoting spacing between groups and reducing direct confrontations that could lead to injury or death. Field experiments in demonstrated that simulated howls often elicited responses from resident packs, such as retreating or remaining stationary, thereby minimizing territorial intrusions without physical encounters. Similarly, in primates like howler monkeys (Alouatta spp.), loud calls analogous to howling deter intergroup incursions and facilitate resource defense, with studies showing heightened vocal activity near group borders. The evolutionary origins of howling trace back to adaptations in early mammals for maintaining group cohesion in low-visibility conditions, such as nocturnal or forested habitats. Fossil evidence from Jurassic mammaliaforms, dating to approximately 165 million years ago, reveals the development of modern hyoid bone structures that support throat functions including swallowing and potentially vocalization. In prehistoric carnivores, hyoid elements from Pleistocene fossils at Rancho La Brea indicate that extinct canids, such as the dire wolf (Canis dirus), possessed robust hyoids capable of supporting howling, suggesting this vocalization evolved as a long-distance communication tool among pack-hunting ancestors. Howling provides key adaptive benefits by enhancing survival and reproductive outcomes through non-aggressive conflict resolution. By signaling presence and intent over distances up to several kilometers, it reduces the energetic and injury costs of physical fights, allowing groups to conserve resources for hunting and pup-rearing. Comparative ethological studies highlight higher reproductive success in howling species; for instance, in captive black howler monkeys (Alouatta caraya), increased howling frequency correlated with greater breeding outcomes, likely due to strengthened pair bonds and reduced stress. Recent research in the employs bioacoustics to explore links between howling traits and genetic fitness. Acoustic analyses of howls have identified individual signatures and pack-specific patterns, enabling noninvasive monitoring that reveals how vocal influences group and rates. For example, studies using on howl spectrograms have shown correlations between acoustic variability and pack cohesion, with more complex choruses associated with larger, fitter groups in wild populations.

Howling in Animals

In Canines

In canines, howling serves as a primary long-distance communication tool, particularly within the family , enabling coordination, territorial defense, and social bonding across species like wolves, domestic dogs, coyotes, and foxes. Among gray wolves (Canis lupus), pack assembly howls facilitate rendezvous after separation, with choruses often initiated by a single individual to gather dispersed members. These howls can be audible over distances exceeding 10 km in open terrain under favorable conditions, such as calm weather and flat landscapes, allowing packs to maintain cohesion over large home ranges. Howling frequency exhibits seasonal variations, peaking in winter during the breeding season when packs are more dispersed and vocal activity supports mate guarding and pair bonding, while rates decline in summer as pups become vulnerable and adults focus on den protection. Studies reveal howl dialects that vary by pack, with distinct acoustic signatures in pitch, duration, and modulation helping to identify group identity and reduce inter-pack conflicts. These dialects emerge from learned behaviors within packs, enabling wolves to recognize allies versus intruders through subtle vocal patterns. Domestic dogs (Canis familiaris) retain as a vestige of their wolf ancestry, though expression varies by due to during . Breeds closer to wolves genetically, such as Siberian Huskies, howl more frequently and respond strongly to wolf-like calls, exhibiting prolonged, melodic vocalizations compared to less ancestral breeds like Labrador Retrievers. Common triggers include high-pitched sounds like sirens, which mimic conspecific howls and elicit instinctive responses for pack coordination or threat alerting, and isolation, where howling signals distress from separation anxiety to summon absent owners or companions. These behaviors underscore the genetic continuity with wolves, as dogs with greater phylogenetic proximity maintain more robust howling repertoires for social signaling. In coyotes (Canis latrans), howling often combines with yips in "group yip-howls," serving as alarm signals to warn packmates of intruders or threats while reinforcing territorial boundaries. These vocalizations proclaim pack presence and size to neighboring groups, deterring encroachment by advertising defensive readiness. Foxes, particularly red foxes (Vulpes vulpes), employ similar yip-howl combinations for alarm and territory defense, with cubs beginning to vocalize, including rudimentary howls, around 3-4 weeks of age as they emerge from dens and integrate into social dynamics. Territorial mapping occurs through howl overlaps, where adjacent canid groups monitor and respond to vocal intrusions, adjusting boundaries based on response intensity and to minimize direct confrontations.

In Primates

Howler monkeys (genus Alouatta), native to Central and South American rainforests, produce some of the loudest vocalizations among land animals, with calls reaching up to 140 decibels and audible from distances of up to 5 kilometers in dense forest environments. These roars, often led by males, form a characteristic dawn chorus that serves primarily to defend territory against intruding groups and deter potential rivals. Male howling intensifies during breeding seasons to signal dominance and attract mates, helping to maintain reproductive access within multi-male troops. The anatomy of howler monkeys is specialized for amplifying these powerful calls, featuring an enlarged that acts as a chamber to deepen and project the sound. This bone, particularly prominent in males where it can be up to four times larger than in females, enables lower-frequency roars that travel farther through the humid, vegetative canopy. Complementing this vocal adaptation is their , which provides stability and mobility in arboreal habitats, allowing individuals to position themselves optimally for howling without compromising locomotion. Sex differences extend to call structure, with males producing longer, more intense roars characterized by deeper pitches, while female calls are shorter and higher in frequency, reflecting dimorphism in hyoid size and throat pouch development. In social contexts, howling reinforces group among of 5 to 20 individuals navigating fragmented arboreal landscapes, where is limited and travel is energy-intensive. These calls coordinate group movements, warn of predators, and facilitate intergroup spacing, reducing physical confrontations in resource-scarce environments like the . Research from the 2010s in Amazonian sites, such as studies on monkeys (Alouatta palliata), has shown that call variation— including roar duration and —correlates with size, with larger groups emitting more synchronized and prolonged choruses to assert presence over broader territories. Among other , (family Hylobatidae) exhibit vocalizations resembling howls in their intensity and territorial function, but these are primarily melodic duets sung by mated pairs to strengthen bonds and advertise occupancy. Unlike the roars of howler monkeys, gibbon songs are structured, species-specific sequences that emphasize pair in small, monogamous family units, distinct in their rhythmic and harmonic qualities.

In Other Species

In felids, howling-like vocalizations are prominent during mating seasons among species such as cougars (Puma concolor) and bobcats (Lynx rufus). Cougars produce intense scream-howls, primarily by females to attract mates, which can carry over distances of up to 1.6 kilometers in open terrain. These calls feature high-pitched, wavering tones that serve as long-range signals in their expansive territories. Bobcats emit similar screaming yowls during courtship, often described as eerie wails that function to locate potential partners across forested areas. Domestic cats (Felis catus) rarely produce true howls in controlled environments but may resort to prolonged yowling or caterwauling in feral conditions, particularly during estrus, mimicking wild felid behaviors to assert territory or seek mates. Among other mammals, certain species exhibit howling analogs adapted to unique ecological niches. Harbor seals (Phoca vitulina) generate underwater roars and moans as pup contact calls, enabling mothers to locate offspring from distances up to 1 kilometer even when separated by water surfaces. These low-frequency vocalizations propagate efficiently through aquatic mediums, facilitating reunions in dynamic coastal habitats. Koalas (Phascolarctos cinereus) produce deep bellows classified as a form of subsonic howl for establishing dominance among males, with frequencies as low as 27 Hz that travel up to several hundred meters through forests during breeding periods. This resonant call, enabled by specialized pharyngeal folds, advertises body size and territorial status without physical confrontation. Avian howling is rare, but some species display convergent vocal traits resembling mammalian howls in function and form, particularly in nocturnal or isolated environments. Certain , such as the (Bubo virginianus), emit deep, rhythmic hoots that echo over wide areas to defend territories or attract mates, akin to howling in their sustained, far-carrying quality. These calls, produced in remote woodlands or islands, illustrate evolutionary parallels where acoustic signaling compensates for visual limitations in dense or dark habitats. Research on howling in non-canine, non-primate species remains limited. For instance, red pandas (Ailurus fulgens) employ hoots and huff-quacks for communication.

Human and Cultural Dimensions

Physiological Capacity in Humans

The human larynx, a cartilaginous structure located in the neck, enables the production of sustained vocal tones through the vibration of vocal folds during phonation, a process facilitated by the coordinated action of laryngeal muscles and airflow from the lungs. This anatomy supports prolonged vocalizations akin to howling, though humans lack the specialized resonance chambers found in some non-human primates, such as the large, hollow bullae on the chimpanzee hyoid bone that enhance low-frequency calls. In humans, the hyoid bone's position and the descended larynx prioritize articulate speech over the resonant, harmonic howls typical of certain primates, resulting in human attempts at howling that are less amplified in lower frequencies. Neurologically, human vocalizations engage the laryngeal responsible for . This shared neural substrate allows humans to imitate animal howls, drawing on limbic influences for emotional intensity similar to those in non-human . Acoustic studies of human vocal imitations reveal that attempts to mimic wolf howls produce sounds with durations standardized to approximately 2 seconds, often exhibiting greater variability and less stability than authentic howls due to the human vocal tract's emphasis on modulation for speech. For instance, human loud vocalizations can reach 80-125 dB, comparable to shouting. In practical applications, therapeutic practices incorporating howling-like vocal releases have emerged in the as part of trends for stress relief, where sustained loud vocalizations activate the to reduce levels and promote emotional . However, prolonged attempts can lead to vocal , including of the vocal folds and , as excessive overtaxes the laryngeal muscles without adequate rest. Such limitations highlight the need for moderation, with recovery typically involving vocal rest to prevent chronic issues like nodules.

Representations in Culture and Media

In , legends frequently portray the creatures as howling under the , symbolizing their transformation and primal fury, with roots traceable to 13th-century sagas like the , where characters don wolfskins and embody lupine behaviors. Similarly, in Native American traditions, wolf spirits are depicted as howling to deliver omens or signal spiritual messages; for instance, the tribe interpreted a wolf's howl as a harbinger of weather changes, while broader Indigenous beliefs across tribes view it as communication from the spirit world. Non-Western perspectives on howling extend to hyena vocalizations in African oral traditions, portraying the animal as a mediator between the living and the dead in cultures from to the . In Japanese folklore, (fox spirits) feature in myths associated with the deity , emphasizing their role as tricksters or messengers. Classical literature references howling to evoke wilderness and foreboding, such as in Virgil's Aeneid (Book 7), where wolves howl amid omens of war and divine unrest, underscoring themes of fate and the untamed natural world. Petronius' Satyricon similarly employs howling imagery in its satirical depictions of excess, with canine howls symbolizing chaos and the blurring of human-animal boundaries during bacchanalian scenes. In 20th-century literature, Jack London's The Call of the Wild (1903) prominently features howling as a primal call to ancestry, as the protagonist Buck joins a wolf pack in a resonant howl that signifies his reversion to wild instincts. In modern media, the howling motif permeates horror films, notably the The Howling series, which began with Joe Dante's 1981 release—a landmark film blending practical effects with on repressed desires, spawning seven sequels that popularized graphic transformation scenes involving howls. During the 2020 , communities worldwide organized nightly "howl-alongs" at 8 p.m. as acts of solidarity, with participants in places like and Mill Valley using simulated howls to honor healthcare workers and express collective resilience amid isolation. In music, traditions incorporate howling as an emotive device, exemplified by artist (Chester Burnett), whose gravelly, howl-infused performances in tracks like "Smokestack Lightning" (1956) capture raw anguish and became iconic in the genre's evolution. Contemporary environmental has repurposed howling simulations in awareness campaigns, fostering public engagement with vocalizations as symbols of ecological balance. Organizations like Howl for Wildlife, active since 2020, advocate for science-based wolf conservation and ethical hunting.

References

  1. [1]
    Wolf Howling Is Mediated by Relationship Quality Rather Than ...
    Aug 22, 2013 · Results showed that relationship quality between the howler and the leaving individual better predicted howling than did the current physiological state.
  2. [2]
    Disentangling canid howls across multiple species and subspecies
    Canid howls are long-range communication signals, used both for territorial defence and group cohesion. Previous studies have shown that howls can encode ...
  3. [3]
    Evolutionary Trade-Off between Vocal Tract and Testes Dimensions ...
    The powerful and characteristic roars of howler monkeys (genus Alouatta) are among the loudest vocalizations produced by any terrestrial animal ...
  4. [4]
    Decoding the Call of the Wild | Science | AAAS
    Aug 22, 2013 · A wolf's howls were a way of reestablishing contact with other pack members after the animals became separated, which often happens during hunts.
  5. [5]
    Wolf species have 'howling dialects' | University of Cambridge
    Feb 8, 2016 · The largest ever study of howling in the 'canid' family of species – which includes wolves, jackals and domestic dogs – has shown that the ...
  6. [6]
    Why Do Dogs Howl? - VCA Animal Hospitals
    Howling is a form of communication that helps members of the pack find each other. Dogs that remain behind vocalize to signal the location of home base and ...
  7. [7]
    Calls of the Wild—Why Do Animals (Including Your Dog) Howl?
    Apr 25, 2015 · All species of howler monkeys howl, especially males: Their hyoid bone, "which serves as a resonating chamber," is larger than those in females.
  8. [8]
    Big testicles mean a softer voice, at least in howler monkeys - Science
    Oct 22, 2015 · Testes size is inversely correlated with animals' ability to roar. The roars of male howler monkeys can be heard up to 5 kilometers away.
  9. [9]
    Howler Monkeys Trade Testicles for Decibels - Scientific American
    Nov 13, 2015 · Biologists have long suspected that the monkeys' howls played a role in attracting mates. The sounds are like the auditory version of ...
  10. [10]
    [PDF] Howling on the edge: Mantled howler monkey (Alouatta palliata ...
    Apr 22, 2019 · Howler monkeys may howl to defend resources such as feeding trees or areas of rich vegeta‐ tion from other monkey groups. This study tests the ...
  11. [11]
    Howling Activity of Mantled Howler Monkeys | Biology
    The howlers call for longer periods of time around 5:00 AM. They also call more frequently in the morning then they do in the afternoon and evening.
  12. [12]
    [PDF] Disentangling canid howls across multiple species and subspecies
    A few studies have examined frequency modulation in canid howls, e.g. in coyotes (Hallberg, 2007) and Iberian wolves (Palacios et al., 2007), by defining ...
  13. [13]
    Genetic distance from wolves affects family dogs' reactions towards ...
    Feb 6, 2023 · Our results suggest that domestication impacts vocal behaviour significantly: disintegrating howling, a central, species-specific communication form of canids.
  14. [14]
    Triggers and consequences of wolf (Canis lupus) howling in ...
    Oct 28, 2022 · In our study, half the howl events were triggered by 12 different environmental or social situations, most of which generated levels of anxiety.
  15. [15]
    'Howl' To Communicate With Your Dog | VMBS News
    Mar 1, 2024 · Howling, on the other hand, is a distinctive and long vocalization, often transitioning from a low to high pitch, that reflects their long-range ...
  16. [16]
    Barking vs. Howling: 6 Reasons Your Dog Howls - Neater Pets
    Mar 8, 2023 · A bark is quick and explosive, while a howl delivers a more sustained vocalization. But what are the differences between barking and howling?
  17. [17]
    Lions' and tigers' fearsome roars are due to their unusual vocal cords
    Nov 2, 2011 · The new study's key finding is that lions and tigers can roar loudly and deeply because their vocal folds have a flat, square shape and can withstand strong ...
  18. [18]
    Howl - Etymology, Origin & Meaning
    Originating in the early 13th century, "howl" is likely imitative, akin to Germanic words; it means to emit a loud, prolonged cry and also functions as a ...
  19. [19]
    Body size and vocalization in primates and carnivores - Nature
    Jan 24, 2017 · DF and F0 were approximately the same (i.e., DF/F0 ≈ 1) in 55% of the primate vocalizations and 36% of the carnivore vocalizations. When DF and ...
  20. [20]
    Decoding Group Vocalizations: The Acoustic Energy Distribution of ...
    May 4, 2016 · We analysed 110 Iberian wolf chorus howls with known pack composition and found that the acoustic energy distribution is concentrated at higher ...
  21. [21]
    1 Dominant frequencies for the exhaling phase of howler monkey ...
    Specifi cally, although all species have their low- est emphasized bands in the 200-700 Hz range, A. caraya and A. palliata produce some of the lowest peak ...
  22. [22]
    Identifying unknown Indian wolves by their distinctive howls - Nature
    Mar 31, 2021 · The average fundamental frequency and duration of Indian wolf howls are 422 Hz and 5.21 s, respectively. Due to its low-frequency range and ...Missing: characteristics | Show results with:characteristics
  23. [23]
    Wolves and Howling - Wolf Conservation Center
    Feb 25, 2017 · Wolves can call to one another over great distances by howling. A howl's low pitch and long duration is well suited for transmission on the wild landscape.<|separator|>
  24. [24]
    [PDF] WOLF HOWLING AND ITS ROLE IN TERRITORY MAINTENANCE
    On all three occasions, weather conditions were optimal, with calm air. Under similar conditions, the maximum distance we heard wolf howling was. 5 km. Thus ...
  25. [25]
    Visualizing sound: counting wolves by using a spectral view of the ...
    Sep 15, 2015 · Wolf howling monitoring technique could be improved by recording wolves' replies and by using bioacoustic tools such as spectrograms and ...
  26. [26]
    [PDF] Acoustic Identification of Wild Gray Wolves, Canis lupus, Using Low ...
    The characteristics of wolf howls such as their maximum frequency, amplitude, duration, and harmonics are used to identify individuals.7, 8 A prior study on ...
  27. [27]
    The Neural Control of Vocalization in Mammals: A Review
    The review describes a model of vocal control, based mainly on research in the squirrel monkey, which consists of two hierarchically organized pathways.
  28. [28]
    Howler Monkey - an overview | ScienceDirect Topics
    Howler monkeys are well known for their loud calls, generated by resonance of the enlarged hyoid bone within a large chamber enclosed by very deep mandibles ( ...
  29. [29]
    Central Nervous System Control of Voice and Swallowing - PMC
    Both vocalization and swallowing are systems evident at birth based on initial brainstem or midbrain control with cortical control for volitional control ...<|control11|><|separator|>
  30. [30]
    Responses of limbic, midbrain and brainstem structures to ...
    The objective of this chapter is to describe the rat limbic, midbrain and brainstem regions that participate in production of vocalizations elicited by midbrain ...
  31. [31]
    Effects of Angle of Epiglottis on Aerodynamic and Acoustic ... - PubMed
    Conclusions: The Aepi was proposed for the first time in this study. The Aepi plays an important role in phonation and resonance of excised canine larynges.Missing: howling mechanism paper
  32. [32]
    Mammalian laryngseal air sacs add variability to the vocal tract ... - NIH
    The mammalian voice apparatus is mainly composed of vocal folds, which work as a sound source generator, and a vocal tract, which works as an acoustic filter.Missing: variations throat
  33. [33]
    The functions of laryngeal air sacs in primates: a new hypothesis
    Laryngeal air sacs in apes and larger gibbons may allow re-breathing exhaled air, enabling fast, extended calls without hyperventilation.
  34. [34]
    Laryngeal Paralysis in Dogs - Veterinary Partner - VIN
    Jun 19, 2006 · Laryngeal paralysis results when the abductor muscles of the larynx cannot work properly. This means no expanding and opening of the larynx for a deep breath.
  35. [35]
    Laryngeal Paralysis - American College of Veterinary Surgeons
    In dogs with laryngeal paralysis, this slowing down is possibly related to poor airflow into their lungs through a compromised larynx. Many families report that ...
  36. [36]
    Wolf Ecology and Prey Relationships on Isle Royale (Chapter 2)
    Distance Traveled by Wolf Packs ... In addition to possible territorial significance, howling helps to assemble individuals in a pack after they have been ...
  37. [37]
    Behavioral and ecological implications of seasonal variation in the ...
    Apr 22, 2017 · Our objective was to examine seasonal changes in the frequency of occurrence of wolf howls and howling replies to understand better their ...
  38. [38]
    [PDF] Environmental and social factors influencing wolf (Canis lupus ...
    Apr 29, 2020 · Howling by gray wolves can also vary with environmental conditions such as season and reproductive activity, time of day, wind conditions, and ...
  39. [39]
    Genetic distance from wolves affects family dogs' reactions towards ...
    Feb 6, 2023 · Of the tested dogs, 57.3% (39 of 68 individuals) reacted with vocalisations in response to wolf howls (for definitions of call types, see ...
  40. [40]
    Do domesticated dogs howl back at wolves? - Popular Science
    Feb 6, 2023 · The dogs more closely related to wolves genetically, such as the Siberian husky, respond to wolf howls more than dog breeds like labrador retrievers.
  41. [41]
    Howling | ASPCA
    Howling is one of many forms of vocal communication used by dogs. Dogs howl to attract attention, to make contact with others and to announce their presence.Missing: scientific definition
  42. [42]
    Probing Question: Why do coyotes howl? | Penn State University
    Jan 14, 2007 · A second reason that coyotes howl is to advertise their presence to other packs, essentially warning those other family groups against ...
  43. [43]
    Structure and Function of Red Fox Vulpes Vulpes Vocalisations
    Aug 7, 2025 · ... barking howls, combination howls, yip howls, whimpers, growls, barks, and choruses. The flat howl of red wolves and coyotes was compared ...Missing: onset | Show results with:onset
  44. [44]
    Red Fox Breeding - Growth & Development of Cubs | Wildlife Online
    The cubs emerge from the natal earth at about five weeks old and may be seen playing outside in late April or early May; for the next few weeks they will ...
  45. [45]
    Black Howler Monkey - Los Angeles Zoo and Botanical Gardens
    ... howler monkeys are considered the loudest land animal in the world. Their calls can be heard up to three miles away and can reach 140 decibels. (A jet ...
  46. [46]
    Black howler monkey - National Zoo
    The large hyoid bone (Adam's apple) that allows for their loud call restricts arm movement, so howler monkeys rely heavily on their tail for locomotion.
  47. [47]
    Alouatta belzebul (red-handed howler monkey) - Animal Diversity Web
    Red-handed howler monkeys males often howl, allowing them to asses their ... Red-handed howler monkeys breed throughout the year and have one offspring ...
  48. [48]
    Evolutionary Trade-Off between Vocal Tract and Testes Dimensions ...
    Oct 22, 2015 · As a result of their anatomical modifications, howler monkeys produce exceptionally low-frequency vocalizations for their body size compared ...
  49. [49]
    Black Howler Monkey - Phoenix Zoo
    Males have much larger hyoid bones and thus make louder calls than females. ... A howler monkey's tail is prehensile (grasping) and acts like another arm ...
  50. [50]
    The function of loud calls in black howler monkeys (Alouatta pigra)
    Our findings suggest that loud calls in black howler monkeys are multifunctional, but most frequently occur in the defense of major feeding sites.
  51. [51]
  52. [52]
    Vocalizations - Gibbon SSP
    This call is also a mechanism for social bonding. Zoos with multiple gibbon groups will often observe gibbons counter-calling. The start of one group's call ...
  53. [53]
    Gibbons singing - Earth.fm
    How do gibbons sing? · 'Black Howler Monkey at Night' · 'Dawn in Khao Yai' · 'Madagascar Dry Forest Sounds' · 'Orangutan Calling' · 'Rincón de la Vieja Dusk' · ' ...
  54. [54]
    Hear a Mountain Lion's Powerful Scream—A Sound You Hope to ...
    Jul 22, 2025 · Their unique screaming behavior occurs during mating season, with females being the primary vocalists.
  55. [55]
    Learn about bobcats | Mass.gov
    Males maintain larger home ranges than females and individuals can travel up to four miles daily. Both sexes use scent marking to mark trails and den sites.
  56. [56]
    The Cat's Meow! Caterwauling in Cats | VCA Animal Hospitals
    Why do cats howl? Cats caterwaul to communicate many needs and emotions, including the following: Physical problems: Cats that are in pain will often make noise ...Missing: domestic | Show results with:domestic
  57. [57]
    Fun Facts About Surprising Seals | NOAA Fisheries
    Aug 20, 2025 · Harbor seals are an exception, and mothers will leave pups on shore to feed offshore. Seal mothers and pups can stay in contact by sound. Harbor ...Missing: howls | Show results with:howls
  58. [58]
    Loud Koalas - California Academy of Sciences
    Dec 19, 2013 · During breeding season, male koalas produce low-pitched 'bellow' vocalizations that are about 20 times lower than would be expected for an ...
  59. [59]
    Koalas' bellows boast about size - ScienceDaily
    Oct 1, 2011 · So, koala males are able to communicate their size, with the largest animals producing the richest baritone bellows. Charlton also suspects that ...
  60. [60]
    Great Horned Owl Sounds, All About Birds, Cornell Lab of Ornithology
    Great Horned Owls advertise their territories with deep, soft hoots with a stuttering rhythm: hoo-h'HOO-hoo-hoo. The male and female of a breeding pair may ...
  61. [61]
    Know Your Nocturnal Neighbors: Nine Owl Calls to Listen For
    Sep 30, 2021 · Eastern Screech-Owls have two main calls. One is a descending, almost horse-like whinny, used to defend territories. The other is an extended ...
  62. [62]
    Endangered Red Panda Facts - SAFE Worldwide
    The red panda has at least seven different vocalizations, including growls, barks, squeals, hoots, bleats, grunts, and twitters.
  63. [63]
    Red Pandas, Climate Change, and the Fight to Save Forests
    Climate change is impacting species across the globe and red pandas—with less than 10,000 left in the wild—are not immune. Average temperatures in Sikkim are ...Missing: vocalizations howling post- 2020
  64. [64]
    Larynx (Voice Box): Anatomy & Function - Cleveland Clinic
    Your larynx is a hollow tube in the middle of your neck, just above your trachea (windpipe) and esophagus. It makes it possible for you to make sounds.
  65. [65]
    [PDF] Air sacs and vocal fold vibration: Implications for evolution of speech
    Comparison of a human and chimpanzee (female) hyoid bone. Note the large bulla of the chimpanzee hyoid. This is hollow on the inside and contains the ...<|separator|>
  66. [66]
    [PDF] The Morphological Evolution of the Vocal Apparatus in the Human ...
    Descent of the hyoid bone, while virtually identical in human and chimpanzee infants, also varies significantly between adult specimens of each species ...
  67. [67]
    Archive: Secrets of Human Speech Uncovered | UC San Francisco
    Feb 20, 2013 · A team of researchers at UC San Francisco has uncovered the neurological basis of speech motor control, the complex coordinated activity of tiny brain regions.
  68. [68]
    Insights into the Neural and Genetic Basis of Vocal Communication
    Studies in humans and a small number of other species have provided insights into the neural and genetic basis for learned vocal communication and are helping ...Introduction · Cortex · Basal Ganglia
  69. [69]
    Vocalization, Neural Mechanisms of - MIT Press Direct
    The tendency for vocalization and human speech to be strongly affected by emotions may suggest that all vocalizations rely at least in part on the ACG-PAG.
  70. [70]
    Humans Mimicking Animals: A Cortical Hierarchy for Human Vocal ...
    Jun 6, 2012 · We found a cortical hierarchy in humans optimized for processing meaningful conspecific utterances. This left-lateralized hierarchy originated near primary ...
  71. [71]
    At How Many Decibels Does A Human Speak Normally
    A normal voice level is between 60-70 dB · A raised voice is between 65-75 dB · A very loud voice is between 75-85 dB · A shouting voice is above 85 dB.
  72. [72]
    Wolves howl like humans, new voice recognition study shows
    Jul 24, 2013 · Holly Root-Gutteridge of Nottingham Trent University has developed sound analysis code that can tell which wolf is howling with 100% accuracy.
  73. [73]
    Scream therapy: howling into the void cured my stress - The Times
    Oct 28, 2023 · It activates the brain's limbic system, which in turn releases cortisol and adrenaline. Screaming physically can help relieve muscle tension. ...Missing: humans relief<|separator|>
  74. [74]
    What You Should Know About Vocal Strain | Mount Sinai Today
    Apr 28, 2025 · Vocal strain can often be treated by resting the voice and speaking at a lower volume for a few days, which will help prevent significant damage ...
  75. [75]
    Vocal Cord Strain – Symptoms and Causes | Penn Medicine
    Vocal cord strain occurs when the muscles and vocal cords are injured through overuse. It can also be a side effect of laryngitis or surgery to treat cancer.
  76. [76]
    The history of werewolves - Historia Magazine
    Oct 5, 2023 · The Völsunga Saga, written down in the 13th century but undoubtedly much older, is one such tale. It features a father and son who don wolfskins ...
  77. [77]
    Wolves in Native American Culture - Wolf Song of Alaska
    Chippewa myths tell of wolves supplying humans with food and hides. The Delaware tribe thought a change in weather might be announced through a wolf's howl. The ...
  78. [78]
    Wolf Myths and Folklore From Around the World
    Feb 20, 2020 · This might be why howling is often considered a message from the spirit world in Indigenous belief. One Nuu-chah-nulth story from Vancouver ...
  79. [79]
    Hyenas in African Lore: Villains or Shamans? - NewsBreak
    Jul 20, 2025 · Among some diviners in Southern Africa, dreaming of a hyena can indicate spiritual calling or warn of enemies using witchcraft. The tracks or ...<|separator|>
  80. [80]
    Kitsune | Meaning, Japanese, Fox, Powers, Mythology, Yokai, & Types
    Kitsune, trickster foxes from traditional Japanese folklore. They are a type of yōkai, a class of supernatural creatures with godlike powers.
  81. [81]
    3.2 Pietas vs furor - Introducing Virgil's Aeneid - The Open University
    Throughout the poem, Virgil uses imagery of powerful forces of nature to describe furor; for example, fire, storm winds and darkness, and lashing waves.Missing: howling Petronius Satyricon
  82. [82]
    The Correct Order To Watch The Howling Movies - SlashFilm
    May 26, 2024 · Joe Dante's 1981 werewolf flick "The Howling" boasts one of the most impressive werewolf transformation scenes in the genre's history.
  83. [83]
    The howling: Americans - let it out from depths of pandemic
    Apr 10, 2020 · Americans are taking a moment each night at 8 pm to howl in a quickly spreading ritual that has become a wrenching response of a society cut off from one ...Missing: alongs | Show results with:alongs
  84. [84]
    Blues | Definition, Artists, History, Characteristics, Types ... - Britannica
    Oct 25, 2025 · The blues is a form of secular folk music created by African Americans in the early 20th century, originally in the South.
  85. [85]
    What We Do - Howl For Wildlife
    We educate the public, both non-hunting and hunting, about the cultural, economic, ecological, conservation, and sustenance sides of hunting.Missing: simulated post- 2020