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Outer ear

The outer ear, also known as the external ear, is the visible, outermost portion of the , comprising the auricle (or pinna) and the external auditory canal, which collectively capture and channel sound waves toward the tympanic membrane to initiate the hearing process. This structure enhances sound detection by amplifying frequencies, particularly those around 3 kHz vital for speech comprehension, and provides directional cues for through its unique shape. The auricle is an elastic cartilaginous framework covered by and skin containing sebaceous and sweat glands, forming an irregular, concave structure attached to the side of the head by ligaments and intrinsic muscles. Its key anatomical features include the , the prominent outer rim that curves around the ; the , a Y-shaped ridge parallel and anterior to the helix that defines the superior and inferior crura; the tragus, a small cartilaginous flap anterior to the canal entrance; the , opposite the tragus; the , a deep fossa at the base leading to the canal; the scapha, a narrow depression between the helix and antihelix; and the lobule, the soft, pendulous lower portion lacking . These components create a funnel-like shape that filters and directs airborne sound vibrations efficiently. The external auditory canal, approximately 2.5 cm long and S-shaped, extends from the concha to the tympanic membrane, lined with , hair follicles, and ceruminous glands that produce protective (cerumen) to trap debris and prevent . Laterally cartilaginous and medially bony, it resonates sound waves, boosting pressure at the by 10 to 15 decibels for frequencies between 2 and 5 kHz, thereby improving auditory sensitivity to environmental noises and human voices. Overall, the outer ear's design not only protects the middle and but also plays a critical role in spatial hearing and acoustic amplification.

Anatomy

Auricle

The auricle, also known as the pinna, is the visible external portion of the outer ear, primarily composed of elastic covered by a thin layer of that includes directly overlying the . This lacks significant subcutaneous fat, allowing the to closely adhere to the underlying for a defined contour. The fibroelastic nature of the provides flexibility while maintaining the auricle's intricate shape through its network of elastic fibers. Key anatomical features of the auricle include the , which forms the prominent outer rim curving superiorly and posteriorly; the , a Y-shaped inner parallel to the with superior and inferior crura that divides the auricle into upper and lower regions; the scapha, a narrow depression between the and ; and the tragus, a small cartilaginous flap projecting anteriorly over the entrance to the external auditory . Additional components are the , a opposite the tragus at the inferior edge of the ; the , a deep bowl-shaped depression adjacent to the external auditory ; and the lobule, the soft, pendulous lower portion lacking and consisting mainly of areolar and . These elements collectively create the auricle's convoluted surface, which is continuous with the surrounding and face via attachments. The auricle exhibits notable variations in size and shape across populations, influenced by genetic and environmental factors, with fibroelastic playing a key role in preserving its form despite these differences. Common shape variations include (most prevalent), followed by triangular, rectangular, and forms. Anthropometric data indicate average adult dimensions of 6-7 cm in height (from superior to inferior lobule) and 3-4 cm in width (maximum breadth across the ), though these metrics show slight differences by and —for instance, longer averages in some North populations compared to others.

External auditory meatus

The external auditory meatus, also known as the external auditory canal, is a tubular passageway that extends from the auricle to the tympanic membrane, serving as the conduit for sound waves into the . It originates from the anteriormost portion of the auricle's and follows an S-shaped (sigmoid) trajectory through the . The canal measures approximately 2.5 cm in length and has an average diameter of 7 to 8 mm, with a notable narrowing at the located in the bony portion. Structurally, the outer one-third of the is cartilaginous, formed by an extension of the auricle's and covered by a thicker layer of that includes follicles, sebaceous glands, and ceruminous glands. The inner two-thirds is bony, composed of the tympanic and squamous portions of the , with thinner, more tightly adherent . The medial end of the canal is sealed by the tympanic membrane, which forms a slight inward projection. The lining of the consists of that supports protective features, including the production of cerumen () by ceruminous and sebaceous glands, which combines with shed cells and trapped to form a waxy barrier. Hair follicles, primarily in the cartilaginous outer portion, further aid in trapping . A protective is the self-cleaning process driven by epithelial migration, where surface cells move outward from the tympanic membrane toward the external opening at a rate of about 0.1 to 0.15 mm per day, carrying cerumen and away to prevent accumulation.

Auricular muscles

The auricular muscles are a group of small skeletal muscles associated with the auricle of the outer ear, divided into intrinsic and extrinsic categories based on their attachments. These muscles overlay the auricular cartilage and are innervated by branches of the (cranial nerve VII). In humans, they are largely vestigial, exhibiting significant atrophy compared to those in other mammals and ancestral humans, with minimal capacity for voluntary movement. The intrinsic auricular muscles connect different regions of the auricle itself, potentially altering its shape. The anterior group includes the helicis major, which originates from the spine of the helix and inserts into the anterior ligament of the helix; the helicis minor, arising from the crus of the helix and inserting into the helix groove; the tragicus, originating and inserting within the tragus; and the antitragicus, which spans the antihelix to the antitragus. The posterior group comprises the obliquus auriculae, extending from the concha to the posterosuperior auricle, and the transversus auriculae, running horizontally across the concha. These muscles are supplied by temporal and auricular-occipital branches of the facial nerve. The extrinsic auricular muscles link the auricle to surrounding structures such as the and . The originates from the and inserts superiorly into the auricle via a , drawing the ear upward; the arises from the same aponeurosis and inserts into the anterior , pulling the ear forward; and the originates from the mastoid process of the and inserts into the posteroinferior , retracting the ear backward. All are innervated by the , with the superior and anterior by its temporal branch and the posterior by its posterior auricular branch. In humans, the auricular muscles serve no essential functional role and are capable of only slight, rudimentary movements, such as a minor "ear wiggle" in individuals with retained voluntary control, reflecting their evolutionary reduction and atrophy relative to more mobile ears in primates and early hominids.

Vasculature and innervation

The arterial supply to the outer ear arises mainly from the posterior auricular artery, a direct branch of the external carotid artery that emerges within the parotid gland and ascends posteriorly to perfuse the majority of the auricle via small auricular branches, as well as the posterior scalp and surrounding structures. This artery forms anastomoses with the anterior auricular branches, which originate from the superficial temporal artery (another external carotid derivative), ensuring comprehensive perfusion of the anterior auricle, including regions toward the tragus and helix. Venous drainage of the outer ear follows a parallel course, primarily through the posterior auricular vein, which collects from the auricle and converges with the posterior of the retromandibular vein to form the , ultimately returning deoxygenated to the . These superficial s are notably susceptible to in the context of regional infections, such as those involving the auricle or external auditory meatus, due to their proximity to potential sites of and bacterial spread. Sensory innervation of the outer ear is predominantly supplied by branches of the and , reflecting its somatosensory role. The , derived from the anterior rami of C2 and C3, provides sensory input to the lower two-thirds of the auricle, including the , , lobule, and , as well as the skin over the angle of the . The , a branch of the mandibular division (V3) of the , innervates the anterosuperior aspects, such as the tragus, crus of the , and external auditory . The , also from C2 (with contributions from C3), supplies the superoposterior auricle and adjacent scalp. Motor innervation to the intrinsic and extrinsic auricular muscles is provided exclusively by posterior auricular and other temporal branches of the (cranial nerve VII). Lymphatic drainage from the outer ear varies by region but generally flows to superficial nodes before deeper chains. Anterior portions, including the tragus and anterior , drain to the parotid and periparotid nodes; posterior aspects drain to mastoid and level V (posterior triangle) nodes; and inferior regions, such as the lobule, route to superficial nodes along the . This multidirectional pattern supports efficient immune surveillance for the exposed auricular tissues.

Development

Embryological origins

The outer ear, or auricle, begins to form at the end of the fourth week of embryonic development from six mesenchymal hillocks known as the hillocks of His, which arise around the dorsal aspect of the first pharyngeal cleft. These hillocks are derived from cells and of the first and second branchial (pharyngeal) arches, marking the initial stage of auricular . By the sixth to eighth weeks of , the hillocks fuse and differentiate into the mature auricular structures: the three hillocks from the first contribute to the tragus, helical crus, and portion of the , while those from the second arch form the , remainder of the , and lobule. This process establishes the basic topography of the auricle, with incomplete merging often occurring at sites such as the pretragal or postauricular regions, predisposing to certain developmental variations. The cartilaginous framework of the auricle develops from condensations of surrounding , which chondrify and shape the by around 20-22 weeks of . Concurrently, the developing auricle undergoes a posterior of approximately 90 degrees relative to the head, aligning it with the lateral aspect of the by the tenth week. The external auditory meatus originates from the first pharyngeal cleft, where an initial ectodermal pit forms at the sixth week and progressively deepens into a tubular canal through epithelial proliferation and canalization, reaching its mature length by the third month of gestation. Genetic regulation plays a critical role in outer ear embryogenesis, with homeobox (HOX) genes such as HOXA1 and HOXA2 directing the patterning and differentiation of branchial arch derivatives, including the auricular hillocks. Disruptions in these genes can affect mesenchymal migration and hillock fusion, highlighting their influence on malformation-prone sites. The outer ear shares embryological continuity with the middle ear ossicles, which also derive from the first and second branchial arches.

Postnatal changes

Following birth, the outer ear undergoes significant growth and maturation, building on its embryological . The auricle, or pinna, exhibits rapid postnatal , reaching approximately 80% of its adult length by age 6 years, with width maturing earlier around 6-7 years while length continues to elongate until 12-13 years. This growth occurs at a faster rate than the overall head, resulting in proportional changes where the auricle becomes relatively larger in adulthood compared to infancy. The external auditory meatus also widens gradually postnatally, with its bony portion forming between 12 and 15 months and achieving full adult configuration by about 9 years of age. Throughout adulthood and into old age, the auricle continues to elongate at an average rate of approximately 0.22 mm per year in length, primarily due to softening and stretching of cartilaginous tissues. Sexual dimorphism in outer ear size becomes evident post-puberty, with male auricles generally larger than female ones, influenced by hormonal factors during adolescence. In later life, age-related changes include stiffening of the auricular cartilage through calcification, which reduces flexibility. Additionally, cerumen production dynamics shift in the elderly, leading to drier wax and higher rates of impaction due to altered glandular activity and reduced self-cleaning of the meatus.

Function

Sound collection and amplification

The auricle, commonly known as the pinna, serves as a parabolic collector that funnels incoming sound waves into the external auditory meatus, concentrating acoustic energy and directing it toward the tympanic membrane for more efficient transmission. This funneling mechanism enhances the collection of sound from the environment, particularly from frontal directions, by reflecting and channeling waves into the ear canal. The convoluted shape of the pinna, including its ridges and folds, contributes to directional filtering that modifies the spectral content of sounds based on their angle of incidence, thereby boosting the overall frequency response at the eardrum. The external auditory meatus functions as an acoustic , amplifying sound pressures through quarter-wavelength , with a in the 2-5 kHz range that is essential for human speech intelligibility. This increases sound pressure levels by approximately 10-15 in this , effectively elevating the intensity of incoming waves before they reach the . The (HRTF), which encapsulates the combined filtering effects of the head, , and outer ear, further characterizes these modifications, introducing direction-dependent peaks and notches—such as enhancements around 6-8 kHz due to pinna interactions—that shape the reaching the . Cerumen, the waxy secretion in the , plays a protective role by trapping dust, , and foreign particles, preventing and maintaining canal lubrication without impeding normal conduction. In physiological amounts, it allows unobstructed wave propagation to the tympanic , ensuring that mechanisms operate effectively.

Sound localization

The outer ear plays a crucial role in by generating and temporal cues that allow the to determine the direction of a sound source in . The pinna and external auditory modify incoming waves through direction-dependent filtering, creating unique acoustic signatures that vary with (horizontal plane) and (vertical plane). These modifications enable precise localization, particularly for broadband sounds, by providing monaural cues and enhancing differences. The convolutions and ridges of the pinna act as a complex acoustic filter, producing frequency-dependent spectral notches and peaks that serve as primary cues for . For instance, in humans, interactions between sound waves and the pinna introduce prominent notches in the 5-10 kHz , with the notch shifting systematically based on the sound's vertical angle—lower frequencies for sounds above the horizon and higher for those below. These pinna-induced spectral cues are essential for distinguishing elevations, as no two directions yield identical spectral profiles at the . Additionally, the pinna aids interaural level differences (ILDs) and interaural time differences (ITDs) through shadowing effects, where the auricle partially obstructs high-frequency sounds (>3 kHz) to the contralateral , amplifying disparities for azimuthal localization. The external auditory further refines these cues by and filtering high frequencies, contributing to azimuth-specific modifications in the overall (HRTF). This , peaking around 2-4 kHz, enhances sensitivity to directional variations in the horizontal plane by amplifying certain spectral components while attenuating others, thus supporting the detection of subtle ILD and ITD shifts. In combination with pinna effects, the ensures that signals carry robust azimuthal information. In humans, the outer ear is particularly vital for vertical localization, where cues like ITD and ILD are minimal, and cues dominate. Listeners rely heavily on pinna-generated notches for resolving front-back and up-down ambiguities, achieving errors as low as 5-10° in free-field conditions with natural head movements. Experimental tests in anechoic environments, using free-field presentations of noise bursts from multiple azimuths and s, confirm that disrupting these outer ear cues—such as by occluding the pinna—severely impairs vertical accuracy, underscoring the structure's specialized role.

Clinical significance

Congenital anomalies

Congenital anomalies of the outer ear encompass a range of developmental malformations present at birth that affect the auricle, external auditory meatus, or surrounding structures, often resulting from disruptions in early embryonic development of the first and second branchial arches. These anomalies can occur in isolation or as part of genetic syndromes, with a reported incidence of approximately 1 in 6,000 to 1 in 7,000 newborns. Unilateral involvement is far more common than bilateral, occurring in approximately 90% of cases for conditions like . Microtia represents the most frequent anomaly, characterized by underdevelopment or hypoplasia of the auricle, ranging from mild structural abnormalities to complete absence (). It affects approximately 1 in 6,000 to 1 in 12,000 births globally, with higher rates in certain populations such as Hispanics and Asians. Aural involves the absence or closure of the external auditory , often co-occurring with in up to 80% of cases, and has a prevalence of 1 in 10,000 to 20,000 births, predominantly unilateral and right-sided. Preauricular tags and pits, benign accessory or depressions anterior to the tragus, occur in about 1 in 12,500 births and may signal underlying syndromic involvement. The etiology of these anomalies is multifactorial, involving genetic and environmental factors such as exposure to teratogens like or during the first trimester. Genetic causes predominate in syndromic cases, including autosomal dominant disorders like (mandibulofacial dysostosis), which features and in nearly all affected individuals due to in the TCOF1 gene. Similarly, (oculo-auriculo-vertebral spectrum) often presents with unilateral , preauricular tags, and epibulbar dermoids, linked to disruptions in cell migration. , caused by in EYA1, SIX1, or SIX5 genes, associates preauricular pits/tags with branchial fistulas and renal in up to 67% of cases. Microtia and aural atresia commonly cause due to malformations of the external canal and associated anomalies, affecting greater than 90% of the involved ears. Microtia/ occurs in isolation in 60-80% of cases. Systemic associations include renal anomalies, such as or , particularly in syndrome (prevalence up to 67%), and derivatives like clefts or fistulas (up to 49%). Preauricular tags/pits carry a modestly elevated of permanent hearing impairment (up to 8 per 1,000 affected infants) and renal issues in syndromic contexts, though isolated cases show rates similar to the general population. Severity of is classified using systems like that of Marx (1926), which grades the auricle as: Grade I (small but vertically elongated with identifiable landmarks), Grade II (vertical remnant lacking landmarks), Grade III (small horizontal remnant or lobule), or Grade IV (). The Weerda classification expands on this embryologic basis, categorizing first-degree (mild with recognizable structures), second-degree (moderate ), and third-degree (severe, peanut-like remnant) anomalies to guide clinical assessment.

Infections and inflammatory conditions

Otitis externa, commonly known as swimmer's ear, is an inflammatory condition of the external auditory canal primarily caused by bacterial infections, most frequently involving and . It often arises from moisture retention in the , which creates an conducive to microbial growth, or from mechanical trauma such as aggressive ear cleaning with cotton swabs. Common symptoms include severe exacerbated by jaw movement or touch, itching, redness, swelling of the canal, and purulent discharge; in severe cases, it can lead to due to canal . Risk factors include frequent swimming, humid environments, and underlying skin conditions like eczema that compromise the canal's protective barrier. The annual incidence of acute otitis externa in the United States is approximately 4 per 1,000 persons, with higher rates among children and active individuals. Treatment typically involves topical antibiotics, such as or drops, combined with ear canal cleaning to remove debris and promote healing; systemic antibiotics are reserved for cases with or immunocompromise. Perichondritis refers to inflammation of the surrounding the auricular , often progressing to and predominantly caused by following , such as ear piercing or burns. Symptoms manifest as painful swelling, , and warmth over the pinna, sparing the lobule due to its lack of , and can rapidly evolve if untreated. Complications include formation requiring , and chronic fibrosis leading to a deformed "" if blood supply to the is disrupted. Prompt administration of anti-pseudomonal antibiotics, such as fluoroquinolones, is essential, often supplemented by incision for purulent collections to prevent permanent deformity. Relapsing polychondritis is a rare autoimmune disorder characterized by recurrent inflammation of cartilaginous structures, including the auricle, due to immune-mediated destruction of proteoglycan-rich tissues. Auricular involvement occurs in up to 90% of cases, presenting with sudden, painful redness and swelling of the ear , typically bilateral and sparing the lobule, which may recur over months or years. The condition arises from autoantibodies targeting and other components, potentially triggered by genetic or environmental factors. Treatment focuses on with corticosteroids for acute flares and disease-modifying agents like for maintenance to mitigate progression to saddle-nose deformity or airway collapse.

Trauma and foreign bodies

Trauma to the outer ear encompasses a range of injuries, primarily resulting from mechanical forces or environmental exposure, which can compromise its structure and function. Lacerations often arise from shearing forces or , leading to partial or complete tears in the auricular skin and . Blunt force , common in contact sports, frequently causes auricular hematomas, where accumulates between the and due to disrupted vessels. If untreated, these hematomas can organize into fibrotic tissue, resulting in the characteristic deformity known as . , caused by prolonged exposure to subfreezing temperatures, affects the exposed auricle by forming ice crystals in tissues, leading to initial numbness followed by blistering and potential tissue damage. The outer ear's rich vascular supply contributes to profuse bleeding in these injuries, necessitating prompt to prevent further complications. Foreign bodies in the outer ear, including and cerumen impaction, can cause obstruction, irritation, or secondary injury. Insects entering the external auditory canal may trigger intense discomfort due to movement, while excessive cerumen buildup leads to impaction, impairing sound conduction and potentially causing . Removal techniques prioritize non-invasive methods initially; irrigation with warm saline is effective for cerumen and small objects, while manual instrumentation using , hooks, or suction is employed for after with oil or lidocaine to halt movement. For live , the process involves first drowning or anesthetizing the creature to facilitate safe extraction under direct visualization. Post-trauma, the outer ear faces heightened risk due to disrupted barriers and bacterial entry, particularly in lacerations or hematomas exposed to contaminants. Avulsion injuries, involving complete or partial detachment of auricular from such as bites or accidents, carry a of if the avulsed segment loses its blood supply, leading to cartilage ischemia and death. The dense innervation of the auricle amplifies sensitivity in these acute events, underscoring the need for immediate evaluation.

Surgical interventions

Surgical interventions for outer ear pathologies primarily address congenital deformities, such as prominent ears and , as well as acquired conditions like and canal stenosis. These procedures aim to restore anatomical form, improve hearing when applicable, and enhance , often performed in staged approaches for complex reconstructions. Techniques have evolved from early 19th-century methods to modern cartilage-preserving and implant-based options, with outcomes emphasizing symmetry, low recurrence, and minimal donor-site morbidity. Otoplasty corrects prominent ears by reshaping auricular to reduce protrusion, commonly involving conchal setback to decrease the concha-mastoid angle. Introduced by in 1968, this technique uses permanent sutures to fix the concha to the mastoid through a posterior incision, often combined with antihelical fold creation via mattress sutures without cartilage excision to preserve flexibility. Historical roots trace to Friedrich Dieffenbach's 1845 procedure, which involved retroauricular skin excision and conchomastoid suturing for posttraumatic prominence. Success rates exceed 90% for achieving symmetrical ear-head distances (typically 16-18 mm), with patient satisfaction around 94% in cartilage-sparing variants; complications include hypertrophic scarring (2-3%), (1-2%), and recurrence (3-5%), managed through early postoperative monitoring. Microtia reconstruction rebuilds the absent or hypoplastic auricle using autologous rib cartilage grafts or alloplastic implants, typically in staged procedures starting at age 6-10 years to allow rib growth. The two-stage Nagata technique harvests ipsilateral 6th-9th costal cartilages to fabricate a three-dimensional framework, followed by lobule transposition and elevation with a costal cartilage block for projection; outcomes yield detailed conchal definition and minimal resorption (less than 5%), though it requires surgical expertise. Alternatively, porous polyethylene (Medpor) implants enable one-stage reconstruction wrapped in temporoparietal fascia, offering rapid results and outpatient feasibility, but with risks of exposure or fracture (1-5%). Recent advances as of 2025 include tissue engineering strategies using 3D bioprinting and stem cells for auricular regeneration, as well as hybrid frameworks combining autologous rib cartilage with porous polyethylene implants to enhance stability and aesthetics. Rib cartilage methods show comparable aesthetic scores to implants, with no material clearly superior; long-term complications include framework resorption (up to 10%), scar contracture, and donor-site pain (5-15%), reported across 29 studies with follow-up exceeding 5 years. Canalplasty addresses congenital aural by creating a new external auditory canal through mastoid drilling, lined with split-thickness skin grafts or canalplasty flaps, guided by the Jahrsdoerfer grading system for candidacy (scores ≥7 predict success). Performed around age 5-6 years, often coordinated with repair, it restores conductive hearing to near-normal levels in 90% of suitable cases. Meatoplasty treats canal stenosis by excising and widening the with advancement flaps or grafts to prevent debris accumulation and infection. Complications encompass tympanic membrane lateralization (25%), restenosis requiring revision (8%), and rare injury (1%), with higher restenosis risk in acquired cases versus congenital.

Evolutionary history

In mammals

In mammals, the outer ear, consisting of the pinna (or auricle) and external auditory canal, represents a key evolutionary innovation that distinguishes them from other vertebrates, enabling enhanced sound collection and environmental adaptation. This structure evolved in mammals (marsupials and placentals), with the pinna serving as a movable flap of and that varies widely in size, shape, and mobility across species. In many mammals, such as and , the pinnae are highly mobile, controlled by auricular muscles that allow independent rotation and elevation up to 180 degrees, facilitating precise by adjusting the ear's orientation toward auditory cues without head movement. In contrast, exhibit reduced pinna mobility due to a shortened ear tip and inward curl of the , limiting active sound directionality and relying more on head movements for localization. The primary functions of the mammalian outer ear include amplifying and directing sound waves while providing physical protection. Mobile pinnae enhance sound directionality by creating interaural intensity differences and cues, improving horizontal and vertical localization accuracy in like , where pinna movements can refine and estimates during prey detection or threat avoidance. The external auditory canal is lined with containing hair follicles, sebaceous glands, and ceruminous glands that produce cerumen (), forming a protective barrier that traps , repels , and prevents debris entry, thus safeguarding the tympanic from and mechanical damage. Specialized adaptations highlight the outer ear's diversity in mammals. In bats, particularly echolocating species from families like and Rhinolophidae, the pinnae exhibit sharp acoustic tuning to the dominant frequencies of their ultrasonic calls (often 30–140 kHz), amplifying echoes for prey detection and Doppler-based flutter analysis through directional gain and interaural differences. Elephants possess exceptionally large pinnae, which primarily function in by facilitating convective heat loss—up to 100% of daily requirements via and flapping, with surface temperatures varying from 14–32°C and heat dissipation of 10–76 W per ear—while incidentally supporting low-frequency hearing through their expansive sound-collecting area. In humans, as a , the pinna is largely immobilized, with vestigial auricular muscles providing minimal movement, yet it retains its funneling role to concentrate sound waves into the , aiding basic directionality despite the loss of mobility.

Comparative anatomy across vertebrates

In fish, there is no external ear structure; instead, sound detection occurs primarily through the , which senses pressure waves via otoliths, and the system, a mechanosensory organ that detects vibrations and water movements along the body surface. This enables fish to perceive low-frequency sounds and nearby disturbances without an outer ear canal or pinna. Amphibians similarly lack an external ear, relying on body conduction for sound transmission to the , with any tympanic membrane, if present, lying flush with the skin surface rather than recessed. In reptiles and birds, the outer ear remains rudimentary, featuring an exposed or superficial tympanic membrane without a protective pinna; reptiles may have a short external auditory in some , but it serves minimal , while possess an oval ear opening directly on the head surface. Middle ear adaptations, such as the single extrastapedial bone in reptiles and , facilitate sound transfer from the tympanum to the , compensating for the absence of external structures. The mammalian outer ear, including the pinna and external auditory , emerged evolutionarily from derivatives of ancient arches, with in the pinna sharing gene regulatory programs repurposed from filaments, as revealed by comparative and gene-editing experiments. This traces back to in , linking pharyngeal development across species. In some aquatic mammals, such as whales, the outer ear has been secondarily lost, with the pinna absent and the external auditory meatus reduced to a vestigial filled with , adapting to underwater hearing via specialized fat pads. The spiracle in certain , a -derived opening behind the eye, represents an early precursor structure in the broader ear , influencing later auditory adaptations.