A sesamoid bone is a small, rounded bone embedded within a tendon or muscle, typically located near a joint where it functions to protect the tendon from excessive pressure, reduce friction, and enhance the mechanical efficiency of muscle action by increasing leverage and redistributing forces.[1] The name derives from the Greek word sēsamoeidēs (σῆσαμοειδής), meaning "sesame-shaped", due to their small size resembling sesame seeds, often just a few millimeters in diameter.[2] These bones are classified as a distinct type of skeletal element, separate from the primary long, short, flat, and irregular bones, and they develop as focal areas of ossification within soft tissues rather than as independent structures.[3]Sesamoid bones occur in various locations throughout the human body, with up to 42 potential sites per individual, though not all are present or ossified in every person.[1] The most prominent and consistently present is the patella, the largest sesamoid bone, which embeds in the quadriceps tendon anterior to the knee joint and articulates with the femur to facilitate knee extension.[1] In the hands, sesamoids are commonly found at the metacarpophalangeal joints of the thumb, index, and little fingers, embedded in the flexor pollicis brevis and adductor pollicis tendons, while the pisiform in the wrist serves a similar role in the flexor carpi ulnaris tendon.[2] In the feet, paired sesamoids—tibial and fibular—are located beneath the head of the first metatarsal in the ball of the foot, within the tendons of the flexor hallucis brevis muscle, where they support weight-bearing and propulsion during gait.[4] Less common variants include the fabella in the lateral gastrocnemius tendon behind the knee and the os peroneum in the peroneus longus tendon of the foot.[1]Functionally, sesamoid bones act like pulleys, altering the direction of tendon pull to optimize muscle force transmission and absorb compressive loads, thereby preventing tendon wear and injury during movement.[2] They are particularly vital in areas of high mechanical stress, such as the knee for knee locking in extension or the forefoot for push-off in walking and running.[4] Developmentally, sesamoids originate as cartilaginous nodules that form during embryonic stages—such as the patella appearing by the ninth week of gestation—and typically ossify during prepuberty or puberty through endochondral ossification, influenced by mechanical forces and genetic factors.[1]Clinically, sesamoid bones are susceptible to disorders including acute fractures from trauma, stress fractures from repetitive loading, and sesamoiditis—an inflammation often seen in athletes due to overuse—which can cause pain, swelling, and impaired joint function.[4] Accessory or variant sesamoids may lead to complications like nerve entrapment or joint instability if symptomatic, highlighting their role in musculoskeletal biomechanics and the need for targeted imaging in diagnosis.[1]
Anatomy
Structure and composition
Sesamoid bones are small, round bones embedded within tendons or joint capsules, often positioned near articulations to facilitate mechanical efficiency, with the patella representing the largest and most prominent example.[1] These bones typically exhibit irregular shapes, ranging from oval to nodular forms, allowing them to conform to the surrounding soft tissues.[5]In terms of composition, sesamoid bones primarily consist of an inner core of trabecular bone surrounded by a thin outer shell of cortical bone, which provides structural support while minimizing weight. The majority of their surface is covered by the enveloping tendon or ligament, except at the articular surfaces where they may interact directly with adjacent bones or form part of a joint.[1] Due to their embedded position, sesamoid bones have a limited blood supply derived from perforating vessels in the surrounding soft tissues, which predisposes them to avascular necrosis under stress or injury.[1] Innervation is similarly provided by nerves from the adjacent tendons, muscles, or joint capsules, varying by anatomical context but generally supporting sensory feedback rather than motor function.[1]Histologically, sesamoid bones feature a dense network of trabeculae within the cancellous interior, populated by a high concentration of osteocytes embedded in lacunae, which maintain the bonematrix through mechanosensory responses.[6] The extracellular matrix is dominated by type I collagen fibers aligned parallel to the principal axes of compressive loading, reinforced by hydroxyapatite crystals to enhance resistance to pressure and shear forces characteristic of their tendinous environment.[6] This orientation optimizes load distribution, reflecting adaptations to the repetitive compressive stresses encountered during movement.[7]
Locations in the human body
Sesamoid bones in the human body are primarily embedded within tendons crossing joints, with the most prominent and consistently present being the patella, located in the quadriceps tendon anterior to the knee joint, serving as the largest sesamoid and universally present in all individuals.[1] In the foot, the two hallmark sesamoids are found at the medial and lateral aspects of the first metatarsophalangeal joint of the hallux (big toe), with a prevalence approaching 100% (99.96%) in large population studies.[8] Smaller sesamoids occur less frequently in other foot locations, such as the lateral aspect of the fifth metatarsophalangeal joint (prevalence of 13%) and occasionally at the interphalangeal joint of the hallux (22.4%).[9]In the upper extremity, sesamoid bones are notably present in the hand, particularly at the thumb's metacarpophalangeal joint, where two sesamoids (medial and lateral) are embedded in the flexor pollicis brevis and adductor pollicis tendons, with a prevalence of nearly 100% in adults.[10] Additional sesamoids appear variably in the hand, including at the thumb interphalangeal joint (~16-42% prevalence), index metacarpophalangeal joint (~34-61%), and little finger metacarpophalangeal joint (~80%), though they are rare in the middle and ring fingers (<2%).[10] In the wrist, the pisiform bone is sometimes classified as a sesamoid due to its position in the flexor carpi ulnaris tendon, though its developmental origin remains debated.[1] At the ankle, an os peroneum sesamoid is located within the peroneus longus tendon near the calcaneocuboid joint, present in approximately 20% of individuals.[5]Accessory sesamoid bones occur with lower frequency and include the fabella, a small sesamoid in the lateral head of the gastrocnemius tendon posterior to the knee, with a global prevalence of approximately 39% (range 3-87%), varying by ethnicity (higher in Asian populations at up to 87%).[11] The cyamella, another rare accessory sesamoid within the popliteus tendon, has an ossified prevalence of 0.57% to 1.8% based on limited cadaveric and radiographic studies.[12]Sesamoid bones generally exhibit bilateral symmetry, with paired occurrences more common than unilateral ones; for instance, fabellae are bilateral in the majority of cases where present.[13]Sexual dimorphism is observed in certain sesamoids, such as the fabella, which is 2.47% to 2.60% more prevalent in males than females, potentially linked to differences in body size and muscle mass.[14] In the hands, some studies indicate females may possess a slightly higher average number of sesamoids, though this variation is not statistically dominant across populations.[15]
Development
Embryological origin
Sesamoid bones originate from mesenchymal precursors embedded within tendons, differentiating into cartilaginous nodules that serve as secondary ossification centers, in contrast to the primary ossification centers that form along the diaphyses of long bones.[1] These precursors are specifically Sox9-positive and scleraxis (Scx)-positive chondroprogenitors located at sites of tendon-bone interaction, such as regions where tendons wrap around joint prominences.[16] This intra-tendinous development distinguishes sesamoids from the intramembranous or primary endochondral processes seen in the main skeletal elements.The initial formation of sesamoid cartilages occurs during the first trimester of human gestation. The patella, the largest sesamoid, begins chondrifying around the 9th week of gestation, becomes fully cartilaginous by the 14th week, and achieves an adult-like structure by the 23rd week.[1] In the foot, sesamoids of the first metatarsophalangeal joint emerge from the joint capsule by the 12th week, with collagenous fiber attachments integrating into their developing cartilage.[17]Genetic and molecular factors, rather than mechanical stress alone, drive this differentiation at the tendon-bone interface. Transforming growth factor-beta (TGFβ) signaling is essential for specifying Sox9+/Scx+ progenitors into sesamoid fates across types like the patella, fabella, and digit sesamoids.[16]Bone morphogenetic protein (BMP) signaling contributes specifically, with BMP4 required for patella differentiation and BMP2 for overall sesamoid maturation. Hox genes also play a role in positional specification, as mutations in genes like Hoxa10 lead to absence or fusion of certain sesamoids, highlighting their involvement in limb patterning that predisposes tendon sites to sesamoid formation.[18] Recent studies in mammalian models indicate that the initial cartilaginous formation of sesamoids proceeds independently of mechanical stimuli, as evidenced by normal development in paralyzed embryos; however, mechanical load is required for subsequent joint cavitation and separation from adjacent bones.[16][1]True sesamoids are defined by their consistent, endochondral development within tendon substance, differentiating them from sesamoid-like structures such as extraskeletal ossifications, which arise outside tendons without tendinous integration, or accessory bones like the os trigonum that form from joint ligaments rather than tendons.[1] This distinction underscores the adaptive, tendon-embedded nature of true sesamoids.
Ossification and growth
Sesamoid bones typically undergo endochondral ossification, beginning as cartilaginous nodules that form within tendons and ossify through the replacement of cartilage by bone tissue.[1] This process initiates in multiple ossification centers during childhood, with the patella—the largest sesamoid—showing initial ossification between ages 3 and 6 years, often starting as multiple foci that coalesce rapidly.[19] In contrast, smaller sesamoids in the foot, such as those at the first metatarsophalangeal joint, begin ossifying later, around ages 8 to 12 years, aligning with pubertal skeletal development.[8]Many sesamoid bones exhibit multipartite development, where ossification occurs from 2 to 4 separate centers, potentially resulting in bipartite, tripartite, or quadripartite forms if the ossicles fail to fuse completely.[20] Fusion typically happens by late adolescence, but incomplete union—seen in up to 33% of hallux sesamoids—can persist into adulthood, influenced by genetic factors and local mechanical stresses during growth.[21]Growth of sesamoid bones is responsive to mechanical loading, adapting their structure according to Wolff's law, which posits that bone remodels in response to applied stresses, leading to increased density and size under tensile or compressive forces from tendon activity.[1] Growth generally ceases around skeletal maturity, between ages 18 and 25, as overall bone elongation and remodeling slow.[1]Bone mineral density in sesamoid bones peaks during young adulthood, reflecting maximal adaptation to loading, but may decline thereafter due to age-related resorption and reduced mechanical stimuli, similar to patterns observed in other trabecular-rich foot bones.[22][23] In the elderly, this can lead to fragmentation or decreased density, increasing susceptibility to stress-related pathology.[5]
Function
Biomechanical roles
Sesamoid bones primarily function as pulleys within tendon sheaths, enhancing the mechanical efficiency of associated muscles by increasing the moment arm and redirecting forces across joints. In the knee, the patella exemplifies this role by altering the direction of the quadriceps tendon force during extension, thereby amplifying the extensor mechanism's leverage by approximately 30-50%.[24][25] Similarly, in the forefoot, the medial and lateral sesamoids under the first metatarsal head serve as pulleys for the flexor hallucis brevis tendon, optimizing the moment arm to improve great toe flexion efficiency during gait and propulsion.[26]These bones also facilitate force distribution by absorbing compressive loads between tendons and underlying skeletal structures, thereby minimizing friction and enabling greater weight-bearing capacity. For instance, the patella transmits quadriceps forces to the tibia while dissipating patellofemoral joint pressures, which can reach 3 to 7 times body weight during deep knee flexion activities like squatting.[1][27] In high-stress regions such as the knee and forefoot, sesamoids redistribute these forces across broader surfaces, reducing localized tendon strain and supporting sustained locomotion.[28]Additionally, sesamoids provide stress shielding to tendons by protecting them from excessive shear and compressive stresses, altering joint reaction forces to prevent tissue overload. This protective mechanism lowers patellofemoral contact pressures and safeguards tendon integrity during repetitive motions, as evidenced by reduced shear exposure in sesamoid-embedded pathways compared to non-sesamoid tendons.[1][5]Finite element analysis (FEA) models have further elucidated these roles, demonstrating how sesamoids optimize load paths in high-demand areas like the knee and forefoot by modulating stress distributions and minimizing peak tendon strains.[29][30]
Protective and adaptive functions
Sesamoid bones play a crucial role in shock absorption within weight-bearingjoints, dissipating impact forces to protect surrounding structures during dynamic activities. In the forefoot, the sesamoids of the hallux function as cushions, distributing ground reaction forces across the first metatarsal head and reducing peak pressures during the propulsion phase of gait. This mechanism helps prevent excessive stress on the metatarsophalangeal joint and underlying soft tissues, particularly in activities involving repetitive loading such as running or jumping.[31][1]These bones also provide essential protection to tendons by acting as barriers against friction and compression where tendons cross bony prominences. The hallux sesamoids, embedded in the flexor hallucis brevis tendon with the flexor hallucis longus passing over them, prevent abrasion during flexion and extension, enhancing tendon durability in areas subjected to repetitive motion like the push-off in walking. By altering the tendon's angle of pull and reducing direct contact with underlying bone, sesamoids minimize wear and potential microtrauma in high-stress regions.[1][32]Sesamoid bones exhibit adaptive remodeling in response to physiological demands, undergoing hypertrophy and increased density under elevated mechanical loads. In athletes engaged in high-impact sports, such as distance running, the sesamoids demonstrate enhanced bone mineral density and trabecular thickening to accommodate greater force transmission, reflecting Wolff's law of bone adaptation.[1][33]Dysfunction or loss of sesamoid bones can lead to pathophysiological consequences, including increased tendon strain and altered biomechanics. Absence or excision of a sesamoid, as seen in certain anatomical variants or surgical interventions, results in heightened tension on the flexor tendons, predisposing them to inflammation or rupture due to unmitigated friction and force redistribution. This vulnerability underscores the sesamoids' integral role in maintaining tendonintegrity under normal loading conditions.[1][34]
Variations
Common anatomical variants
Sesamoid bones exhibit common anatomical variations in number, size, fusion, and presence across different body regions, reflecting normal developmental diversity in healthy populations. These variants are typically asymptomatic and identified incidentally through imaging or dissection studies. Bipartite or tripartite forms, where a sesamoid fails to fully ossify as a single unit, are among the most frequently observed, particularly in the foot.[8]In the hallux, bipartite sesamoids occur in 10-30% of individuals, with the medial sesamoid more commonly affected than the lateral; tripartite forms are rarer but can appear as multiple separate ossicles without clinical symptoms. These partitions result from incomplete fusion during ossification and are often bilateral, occurring in 80-90% of contralateral feet. Population-based radiographic studies confirm high overall presence of hallux sesamoids, with paired medial and lateral bones identified in approximately 99.9% of cases via X-ray, though the medial sesamoid is consistently visualized in over 85% on MRI and CT imaging in asymptomatic cohorts.[32][35][8]The fabella, an extra sesamoid within the lateral head of the gastrocnemius tendon, shows significant ethnic variation in prevalence, ranging from 10-20% overall but reaching 30-92% in Asian populations and as low as 3-31% in Caucasians. This variation is attributed to genetic and evolutionary factors, with higher rates observed in East Asian and Oceanian groups through systematic meta-analyses of knee imaging.[36][37]In the hand, sesamoid bones are constant at the metacarpophalangeal (MCP) joint of the thumb but variable in other fingers, with absence or hypoplasia common in the index (prevalence ~39%), middle (~99%), ring (~99.5%), and little (~41%) fingers at their MCP joints. Bilateral asymmetry, where sesamoid presence or size differs between hands, occurs in approximately 20% of cases, though lower rates of 5-10% are reported for specific finger asymmetries in some cohorts. These hand variants are well-documented in radiographic surveys, highlighting their role as normal anatomic diversity rather than anomalies.[38][15]
Rare and pathological variants
Rare sesamoid bones, distinct from more common variants, occur infrequently and are often located in tendons away from typical sites. The cyamella, a sesamoid bone embedded in the proximal tendon of the popliteus muscle near the knee, has an ossified prevalence of 0.57% to 1.8% in human populations.[12] This structure is more prevalent in animals, such as 100% in cats and 84% in dogs, but remains a rare normal variant in humans.[39]Accessory sesamoids in the upper extremities, such as the tricipital sesamoid in the tricepstendon or os subepicondylare mediale near the medial epicondyle, are exceptionally uncommon and can mimic avulsion fractures on imaging.[40][41]Congenital aplasia of sesamoid bones, or their complete absence, is a rare anomaly often associated with genetic syndromes disrupting limb development. In Holt-Oram syndrome, an autosomal dominant disorder caused by TBX5 gene mutations, upper limb defects including thumb hypoplasia reflect broader radial ray deficiencies.[42] Such absences are not typically reported in foot sesamoids for this syndrome but underscore the genetic basis of sesamoid dysmorphology in skeletal patterning disorders.[43]Acquired pathological changes to sesamoid bones arise from mechanical stress or degenerative processes, altering their morphology over time. In osteoarthritis, sesamoids may exhibit erosion or fragmentation due to cartilage loss and subchondral bone remodeling at tendon-bone interfaces, particularly in the hallux.[5] Among athletes, repetitive loading can induce hypertrophy or reactive ossification in sesamoids like those of the first metatarsophalangeal joint, exacerbating injury risk.[44] The os peroneum, a sesamoid in the peroneus longus tendon near the cuboid, shows variable prevalence from 5% to 30% across populations, with some cases developing pathological hypertrophy or fracture from chronic stress.[45]Distinguishing pathological variants from acute injuries requires careful imaging to avoid misdiagnosis. Radiographs typically show bipartite sesamoids with smooth, well-corticated margins and symmetric fragments that do not perfectly align, contrasting with irregular, displaced edges in acute fractures.[5]Bone scintigraphy or MRI can further differentiate by demonstrating increased uptake in fractures versus normal activity in congenital bipartites, aiding in the identification of erosion or hypertrophy as acquired changes.[46][47]
Clinical significance
Disorders and injuries
Sesamoiditis is an inflammatory condition primarily resulting from repetitive stress or overuse of the sesamoid bones, commonly affecting athletes such as runners and dancers who engage in high-impact activities like soccer, tennis, basketball, and ballet.[48][49] This overuse leads to irritation of the soft tissues surrounding the sesamoids, particularly at the plantar aspect of the first metatarsophalangeal joint (1MTPJ).[48] Symptoms typically include gradual onset of forefoot pain that worsens with weight-bearing activities such as walking or running, along with localized swelling, tenderness to palpation, and pain during passive extension of the 1MTPJ.[50][51]Fractures of sesamoid bones can occur as acute injuries from high-impact trauma or as stress fractures due to chronic repetitive loading, with the latter presenting as gradual unilateral plantar pain, often involving the medial (tibial) sesamoid of the hallux.[52] In the foot, hallux sesamoid fractures account for a small but notable portion of forefoot injuries, while in the knee, patellar sleeve fractures—a type of avulsion injury involving the cartilage sleeve—are rare and primarily affect children aged 8 to 12 years during sudden quadriceps contractions.[53] These fractures are more prevalent in athletic populations.[54]Avascular necrosis (AVN) of sesamoid bones arises from disrupted blood supply, exacerbated by the inherently poor vascularity of these intra-tendinous structures, leading to bone death and collapse.[1] It is more common in females than males, particularly affecting young women aged 18 to 29 who experience mechanical overload from activities involving repetitive toe-off motions.[55] Symptoms manifest as persistent forefoot pain during the toe-off phase of gait, often without acute trauma, and can progress to joint stiffness if untreated.[55] This condition is analogous to Freiberg's disease but specifically targets the sesamoids, with higher incidence in athletic females due to factors like hormonal influences and training intensity.[56]Bipartite sesamoids, an anatomical variant where the bone fails to fuse during development, have a prevalence of 7% to 30% in the general population, with 90% involving the tibial sesamoid and 80% to 90% being bilateral.[57] In athletes, particularly ballet dancers, these multipartite bones are prone to stress injuries from repetitive hyperextension and loading, leading to inflammation or microfractures at the synchondrosis site and contributing to chronic pain mimicking sesamoiditis.[57] Such stress reactions are noted in up to 4.2% of athletic lower limb injuries, with higher rates in dance populations due to the demands of en pointe positions.[49]
Diagnosis and treatment
Diagnosis of sesamoid bone disorders typically begins with a thorough clinical examination, focusing on palpation of the affected area to identify localized tenderness, particularly under the first metatarsal head for hallux sesamoids or around the patella for knee involvement.[58]Gait analysis may reveal antalgic patterns or pain exacerbated by hyperextension of the great toe in foot cases, helping to differentiate sesamoid issues from conditions like turf toe or metatarsalgia.[32]Imaging plays a crucial role in confirming the diagnosis and assessing the extent of injury. Standard anteroposterior and lateral X-rays are initial tests for detecting fractures, though oblique views are often superior for visualizing the sesamoids due to their embedded position within tendons.[5]Magnetic resonance imaging (MRI) is preferred for evaluating soft tissue inflammation, bone marrow edema, and associated tendinopathy, showing high sensitivity for sesamoiditis and stress reactions.[28]Bone scintigraphy can identify early stress fractures or avascular necrosis when X-rays are inconclusive.[59] Computed tomography (CT) provides detailed bony architecture for complex fractures or bipartite sesamoids mimicking pathology.[60]Treatment approaches prioritize conservative measures, which are effective for most cases of sesamoiditis and acute fractures. Initial management includes rest to offload the sesamoid, nonsteroidal anti-inflammatory drugs (NSAIDs) for pain and swelling reduction, ice application, and elevation. Orthotic devices, such as dancer's pads or custom insoles, help redistribute pressure and prevent further irritation, while immobilization with a walking cast or boot for 4-6 weeks supports healing in foot sesamoid injuries.[61] For patellar sesamoid fractures, immobilization in a knee brace or cast for 6-8 weeks promotes union in stable, non-displaced cases.[62]Surgical intervention is reserved for refractory cases, such as nonunions or chronic pain persisting beyond 6-12 months of conservative care. Excision of the damaged sesamoid is the most common procedure for hallux cases, with low complication rates (around 5-6%) and high patient satisfaction (over 90% return to activity).[63] In patellar fractures requiring surgery, tension band wiring or partial patellectomy achieves union rates exceeding 95%, though nonunion complicates about 1-5% of cases.[64] Post-operative rehabilitation focuses on gradual weight-bearing and strengthening to restore function, with full recovery typically within 3-6 months for foot injuries and up to 6 months for patellar cases.[62]
Comparative anatomy
In non-human mammals
In horses, sesamoid bones are prominent in the limbs, particularly the proximal sesamoid bones (PSBs), which are paired small bones located at the palmar aspect of the fetlockjoint on both fore and hind limbs. These bones are embedded within the suspensory ligament and distal sesamoidean ligaments, functioning to anchor the suspensory apparatus, reduce friction on tendons, and facilitate smooth fetlock flexion during locomotion.[65][66] The PSBs bear significant compressive and tensile forces, especially in high-speed activities, making them susceptible to fractures in racing horses due to repetitive stress and overload.[67][68]Among non-human primates, sesamoid bones exhibit variation, with many types present in most families but frequently reduced or absent in apes, such as the great apes (Hominidae), where pollical metacarpophalangeal sesamoids in the thumb region are often lost or underdeveloped compared to more primitive primate conditions.[69][70] In the giant panda (Ailuropoda melanoleuca), a member of the order Carnivora but noted for its specialized forelimb anatomy, the radial sesamoid bone is greatly enlarged and elongated, forming a "false thumb" that protrudes from the wrist to oppose the true digits. This structure, not fully embedded in tendon like typical sesamoids, enables precise grasping and manipulation of bamboo stems during feeding.[71][72]In rodents, sesamoid bones show diversity, including the cyamella, a small sesamoid in the popliteus tendon at the posterolateral knee, which occurs as a normal variant though rarely documented in specific species. Fabella variations are also noted, with the lateral fabella—a sesamoid in the gastrocnemius tendon—present in many rodents but varying in size, ossification, and bilateral symmetry across individuals and populations. In cotton rats (Sigmodon spp.), a radial sesamoid in the manus articulates with the scaphoid or scapholunar bones, serving as a "false thumb" to enhance grasping of food or objects, analogous to adaptations in other mammals.[73][74][75]Carnivores commonly possess the patella as a large sesamoid in the quadriceps tendon at the knee, alongside fabellae—sesamoid bones embedded in the gastrocnemius muscle tendons behind the femoral condyles, typically one in each head (medial and lateral). These structures enhance knee stability and muscle leverage during quadrupedal movement. In elephants (Elephas and Loxodonta spp.), the prepollex sesamoid in the manus functions as a "sixth toe" or false digit, an enlarged ossified sesamoid that supports weight distribution and pillar-like foot posture, aiding in bearing the animal's massive trunk and body mass.[76][77][78]
Evolutionary perspectives
Sesamoid bones trace their origins to early tetrapods, emerging around 300 million years ago during the Carboniferous-Permian transition, as intratendinous or intraligamentous structures that enhanced joint function in the nascent limb skeleton of land vertebrates.[79] These elements likely arose through the detachment and displacement of epiphyseal or apophyseal tissues from long bones, allowing for greater phenotypic flexibility in skeletal morphology across amniote lineages.[79]Fossil evidence from Permian synapsids, such as basal pelycosaurs and therapsids, reveals early sesamoid-like carpals and tarsals that supported forelimb and hindlimb mobility, indicating their role in the adaptive shifts toward more efficient terrestrial locomotion in proto-mammalian ancestors.Throughout vertebrate evolution, sesamoids exhibit patterns of independent gain and loss, reflecting selective pressures tied to locomotor demands rather than strict phylogenetic inheritance. Absent in fish due to the lack of true limbs, they are variably present in reptiles and birds; for instance, the patella—a prominent sesamoid—is absent in non-avian dinosaurs and most crocodilians but evolved in squamates and most birds to optimize knee extension during bipedal or flight-related activities.[80] In mammals, the patella arose independently four to six times within the crown group, often linked to the biomechanical needs of endothermic locomotion, such as improved quadriceps leverage for sustained activity in monotremes, multituberculates, and therians.[80] A striking example of de novo evolution is the giant panda's radial sesamoid, or "false thumb," which enlarged as a neomorph around 20–30 million years ago in Miocene ancestors like Simocyon, adapting from a wrist bone to facilitate bamboo manipulation in a specialized herbivorous niche.[81]The diversification of sesamoids during adaptive radiations is closely associated with ecological lifestyles, where variations in size, position, and number respond to mechanical stresses in cursorial versus arboreal habitats. In cursorial mammals like equids, patellar sesamoids exhibit thickened cortices and oriented trabeculae to withstand high-speed, ground-based forces, enhancing knee stability during extended gaits. Conversely, arboreal species, such as certain primates and bats, feature more variable or enlarged sesamoids in the manus to support grasping and suspension, promoting diversification into tree-dwelling niches. This modern diversity underscores niche specialization, with sesamoid morphology reflecting evolutionary trade-offs in locomotor efficiency across vertebrates. Genetically, these patterns are influenced by limb-patterning genes like HoxA11, whose mutations disrupt sesamoid formation in experimental models, alongside signaling molecules such as BMPs and GDF-5 that integrate mechanical cues with developmental thresholds.1097-0185(19991015)257:5<174::AID-AR6>3.0.CO;2-O)