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Pygostyle

The pygostyle is a fused skeletal structure at the distal end of the vertebral column in birds, formed by the ossification of the final several caudal vertebrae into a compact, triangular or ploughshare-shaped bone.^[1] This element anchors the rectrices, or tail feathers, and the associated musculature, enabling precise control of the tail as a rudder for steering, braking, and stability during flight, particularly in takeoff and landing maneuvers.^[1]^[2] In avian anatomy, the pygostyle's rigidity contrasts with the flexible tails of reptilian ancestors, reflecting adaptations for powered flight where a fan-shaped tail provides aerodynamic surfaces rather than a prehensile or balancing appendage.^[2] The pygostyle's evolutionary origins trace to the transition from long-tailed theropod dinosaurs to short-tailed avialans, with fusion occurring independently in some non-avian maniraptorans but becoming a defining synapomorphy of the Pygostylia clade among crown-group birds and their close relatives.^[2]^[3] Fossil evidence indicates that early birds like Archaeopteryx retained elongate caudal series without fusion, while later Cretaceous forms developed the pygostyle alongside rectricial bulbs—fibroadipose structures supporting feather vanes—for enhanced tail fanning.^[2] Ontogenetically, pygostyle formation involves progressive vertebral fusion post-hatching, with trabecular remodeling that eliminates intervertebral discs while preserving a spinal cord channel, underscoring its role in lightweight skeletal efficiency.^[3] This structure's morphology varies phylogenetically, correlating with tail feather arrangements and ecological niches, such as dorsoventrally stacked rectrices in certain arboreal or foraging birds.^[4]

Anatomy and Morphology

Structure and Composition

The pygostyle forms the terminal element of the avian vertebral column, resulting from the co-ossification of the final 4 to 6 caudal vertebrae into a single, rigid bony structure unique to birds within Pygostylians.^[3] This fusion yields a laterally compressed, ploughshare-shaped ossification, often triangular in outline with a flattened dorsoventral profile that tapers caudally.^[5]^[6] Key anatomical features include a central hemal canal traversing the structure to house the caudal artery, vein, and associated nerves.^[7] Transverse processes, most developed at the cranial margin, project laterally to provide anchorage for tail musculature.^[8] The ventral surface features an expanded area supporting the rectricial bulb, a fibroadipose complex anchoring the proximal calami of rectrices, while the dorsal surface interfaces with feather follicles and overlying integument.^[9]^[8] Empirical assessments in extant species reveal the pygostyle's brevity relative to overall skeletal proportions, with its length comprising approximately 5-15% of femur length—a standard body size surrogate—contrasting the elongate, discrete vertebrae of non-avian theropod tails.^[5] This compact form enhances structural integrity over the modular composition of unfused caudal elements, though specific bone density metrics remain underexplored in comparative studies.^[4]

Variations in Avian Groups

Palaeognaths, including ratites like ostriches, typically exhibit a pygostyle with incomplete fusion of the caudal vertebrae, resulting in a shorter and less rigidly integrated structure compared to neognaths.^[10] Neognaths, by contrast, display a more compact and fully fused pygostyle, particularly evident in small-bodied clades such as passerines, where the structure supports precise maneuverability in perching and flight.^[11] Morphometric analyses reveal correlations between pygostyle morphology and ecological adaptations, such as foraging styles, across avian taxa. Aerial and pursuit-oriented foragers, including certain insectivores and divers, tend to possess elongated pygostyles, facilitating enhanced tail control during dynamic locomotion, whereas ground-dwelling species often feature more robust, shortened forms suited to terrestrial stability.^[11]^[12] These patterns hold without establishing direct causation, as variations may reflect broader flight and behavioral demands rather than singular ecological drivers.^[13] Among extinct avian groups, enantiornithines frequently displayed a plough-shaped pygostyle distinct from the blade-like form typical of ornithuromorphs, the lineage leading to modern birds; however, convergent evolution produced ornithuromorph-like pygostyles in some enantiornithine lineages, such as Cruralispennia, underscoring homoplasy in tail skeletal adaptations.^[14]^[15] This morphological convergence highlights repeated selective pressures on pygostyle form independent of shared ancestry.^[14]

Development and Ontogeny

Embryonic Formation

The embryonic formation of the pygostyle begins with the segmentation of somites from the tail bud in avian embryos, typically initiating around Hamburger-Hamilton stage 20 in model organisms like the domestic chick (Gallus gallus domesticus), where paraxial mesoderm cells undergo sequential budding to form caudal somites that give rise to sclerotomes and subsequent vertebral primordia.^[16] These somites number approximately 10-12 in the caudal region, producing distinct vertebral elements including centra and neural arches through chondrification and early ossification processes driven by a balance of cellular proliferation, apoptosis, and differentiation.^[17] Hox gene expression patterns, particularly the collinear activation of posterior paralogs (Hox9-13), establish regional identity and correlate with the progressive slowdown of axis elongation, limiting the total number of caudal vertebrae compared to ancestral reptilian conditions.^[18]^[2] In chick embryos, the distal caudal vertebrae emerge as unfused, independent structures, with ossification of free caudal centra beginning around embryonic day 18-20 (near hatching at day 21), while the terminal 4-6 elements retain separate morphology as cartilaginous or early bony precursors without ankylosis.^[19] This initial separation sets the developmental stage for postnatal remodeling, as evidenced by histological analyses showing distinct intervertebral discs and no fusion in late-stage embryos.^[5] Comparative embryology reveals that reptile embryos, such as those of lizards or crocodilians, generate a longer series of caudal somites—often exceeding 20-30 vertebrae—due to extended somitogenesis and sustained Hox-mediated tail bud activity, resulting in persistently unfused tails that contrast with the abbreviated avian pattern.^[20]^[2] These differences underscore Hox-regulated cessation of somite addition as a key mechanism in avian tail truncation during embryogenesis.^[18]

Postnatal Fusion and Inflammation

In extant birds, postnatal fusion of the terminal caudal vertebrae into the pygostyle occurs progressively after hatching, typically completing near skeletal maturity and often post-fledging, as demonstrated by micro-computed tomography (micro-CT) imaging of chicken specimens showing initial unfused centra at 8 days post-hatching with distal-to-proximal coalescence over weeks.^[5]^[21] This process involves ossification, remodeling of intervertebral discs, and resorption of nuclei pulposi, delaying full pygostyle consolidation until late ontogeny despite earlier mineralization of individual vertebrae.^[21]^[22] Sterile inflammation, rather than pathological processes, drives this fusion, as evidenced by histological analysis revealing inflammatory markers and necroptosis—a programmed cell death mechanism—in degenerating disc tissue of developing chicken pygostyles.^[21] Experimental suppression of inflammation via corticosteroid administration (e.g., prednisolone) delays disc degeneration, reduces necroptosis-related gene expression (such as RIPK3), and postpones vertebral fusion, confirming inflammation's causal role without inducing disease.^[21]^[22] This non-infectious response facilitates adaptive skeletal remodeling for flight support, distinct from injury-induced healing. Timing varies across avian developmental modes: in precocial species like domestic chickens (Gallus gallus domesticus), fusion initiates shortly post-hatching and advances rapidly alongside early mobility, achieving near-completion by 8 weeks as per micro-CT scans of control groups.^[22] Altricial passerines, such as the Eurasian reed warbler (Acrocephalus scirpaceus), exhibit delayed overall skeletal ossification sequences, implying protracted pygostyle fusion relative to body growth, though species-specific micro-CT data remain limited.^[23] These differences align with ecological demands, where precocial chicks require earlier tail stabilization for ground-based locomotion and fledging.^[21]

Evolutionary History

Precursors in Non-Avian Theropods

In non-avian theropods, particularly within the maniraptoran clade, fossil evidence reveals partial fusion of distal caudal vertebrae, representing convergent precursors to the avian pygostyle rather than direct homologues. These structures provided increased tail rigidity, likely enhancing balance and postural stability in bipedal locomotion by countering gravitational forces on elongated tails, as biomechanical analyses of theropod caudal morphology indicate.^[24] Such fusions predate the full consolidation seen in avialans and appear sporadically across theropod lineages, underscoring independent evolutionary responses to biomechanical demands rather than a linear progression toward flight.^[2]^[25] Beipiaosaurus inexpectus, a basal therizinosauroid from the Early Cretaceous Yixian Formation of Liaoning, China (approximately 125 million years ago), exhibits a pygostyle-like fusion of the five posteriormost caudal vertebrae, characterized by ventrally concave centra and dorsally arched neural arches forming a compact, blade-shaped terminal structure.^[26] This fusion, observed in holotype specimen IVPP V11559, contrasts with the more elongate, unfused tails of earlier theropods and aligns with reduced caudal counts (around 30 vertebrae total) in derived maniraptorans, facilitating a stiffened tail base for maneuverability.^[27] Therizinosauroids like Beipiaosaurus, though herbivorous and not closely related to avialans, demonstrate that distal co-ossification evolved convergently in response to body mass distribution and integumentary loading, independent of aerodynamic specialization.^[2] Oviraptorosaurs, another maniraptoran group from the Yixian Formation and equivalent strata, display tail reduction with partial distal fusions, though less extensive than in birds. For instance, specimens of Caudipteryx and Similicaudipteryx feature fewer than 25 caudal vertebrae, with the terminal 2–4 co-ossified into a short, rigid unit sometimes termed a pseudo-pygostyle, providing anchorage for proximal tail feathers and counterbalance during terrestrial agility.^[3]^[28] In Nomingia, up to four fused terminal vertebrae exhibit widened transverse processes persisting posteriorly, enhancing lateral stability without full pygostyle consolidation.^[28] These traits, documented in multiple Yixian taxa dated to 120–130 million years ago, reflect adaptations for brooding behaviors and rapid turns in cluttered environments, as inferred from associated nest fossils and skeletal proportions, rather than precursors to avian tail fanning.^[2]^[28]

Appearance in Early Avialans and Birds

The pygostyle emerges in the fossil record as a defining feature of the Pygostylia clade within Avialae, marking a transition from the elongated, unfused caudal series observed in Late Jurassic forms like Archaeopteryx lithographica (circa 150 million years ago) to a condensed terminal fusion in Early Cretaceous avialans.^[2] This reduction involved the fusion of the distalmost caudal vertebrae, typically 4-6 elements, into a rigid structure, first evidenced in deposits of the Jehol Biota, northeastern China, dated to approximately 131-120 million years ago.^[29] Earliest definitive pygostylians include Protopteryx fengningensis, an enantiornithine from the 130.7-million-year-old Sichakou Member of the Huajiying Formation, which preserves a short tail ending in a fused pygostyle, establishing the clade's presence by the Barremian stage of the Early Cretaceous.^[30] Similarly, basal taxa such as Sapeornis chaoyangensis from the younger Jiufotang Formation (circa 120 million years ago) display a triangular pygostyle approximately 11-20 mm long, formed by weakly to moderately fused terminal vertebrae, positioning Sapeornis near the base of Pygostylia.^[31] These specimens indicate that pygostyle development preceded diversification into major ornithothoracine lineages, with early ornithothoracines like confuciusornithids also bearing this synapomorphy.^[32] Confuciusornis sanctus, from the approximately 125-million-year-old Yixian Formation, exemplifies incipient pygostyle fusion, where the terminal four caudal vertebrae exhibit partial coalescence, shorter than the preceding unfused elements, highlighting a stepwise morphological progression in tail abbreviation among primitive pygostylians.^[2] This pattern underscores the rapid skeletal miniaturization in early avian evolution during the Early Cretaceous, with fossil evidence from multiple Jehol localities confirming the pygostyle's establishment prior to more derived ornithuromorph configurations.^[29]

Coevolution with Tail Feathers

The reduction of the avian tail skeleton into a pygostyle coincided with the elaboration of rectrices into aerodynamic fans, enabling precise control during flight through coordinated skeletal support and muscular actuation. Analysis of extant birds reveals that pygostyle morphology—such as its length, robustness, and presence of processes—correlates directly with rectrix fan shapes (e.g., forked, graduated, or square), reflecting adaptations to locomotor demands like foraging or aerial maneuvering.^[33]^[34] This pattern supports coevolution, where skeletal shortening compensated for lost vertebral leverage by enhancing muscle anchors for feather deployment. In Early Cretaceous avialans from the Jehol Biota, including Yixian Formation specimens dated to approximately 125–120 million years ago, pygostyle variations paralleled rectrix asymmetry and fan development across clades. Enantiornithines typically featured elongated pygostyles with prominent uncinate processes, accommodating simpler tail plumage like elongated ribbons or uncinate feathers rather than fully fanned arrays.^[9]^[25] In contrast, ornithuromorphs displayed compact, ploughshare-shaped pygostyles, which anchored robust rectricial bulbs and supported asymmetric vanes in fanned rectrices for superior aerodynamic stability.^[14]^[35] These differences, observed in fossils like Pengornis and Yixianornis, indicate clade-specific evolutionary trajectories in tail function prior to the enantiornithine extinction.^[36] Fossil imprints from Yixian ornithuromorphs preserve rectrices emerging from pygostyle-associated soft tissue, evidencing muscle insertion scars that enabled fanning motions analogous to modern bulbi rectricium.^[8] Such preserved morphologies, including vascular impressions on the pygostyle, demonstrate how skeletal compaction facilitated integumentary elaboration without sacrificing control, as corroborated by comparative dissections linking these features to powered flight enhancements.^[3]^[37]

Functional Adaptations

Skeletal Support for Feathers and Muscles

The pygostyle functions as the principal insertion point for avian tail muscles, enabling precise control over tail posture and feather orientation. Specifically, it anchors the musculus levator caudae, which elevates the tail by contracting dorsally, and the musculus depressor caudae, which lowers it via ventral attachment, with these muscles originating from the synsacrum and ilium.^[38]^[39] In pigeons, for instance, these muscles innervate motoneurons that coordinate pygostyle movements, distributing contractile forces across the fused structure to minimize shear stress.^[38] The pygostyle also supports the rectricial bulbs, paired fibroadipose tissues enveloping the bases of the rectrices (tail feathers), where bulbi rectricium muscles insert to erect and fan the feathers.^[9]^[40] This arrangement allows biomechanical leverage for feather deployment, as the bulbs affix directly to the pygostyle's ventral and lateral surfaces, transmitting tension without requiring extensive skeletal elongation.^[9] Fusion of the terminal 4–6 caudal vertebrae into the pygostyle confers rigidity, concentrating muscle forces onto a single, plow-shaped ossification that resists deformation under load, as demonstrated in dissections of species like woodpeckers where expanded ventral surfaces enhance attachment area.^[4]^[1] This consolidation reduces skeletal mass—typically comprising less than 1% of body weight in many birds—compared to flexible, multi-vertebrate tails, while maintaining structural integrity for force distribution, verifiable through comparative avian anatomy where unfused tails demand additional intervertebral stabilizers.^[41]^[42] In galliform birds such as chickens, tail muscle myology reveals depressor caudae and levator caudae as the dominant components, with masses ranging 0.028–0.329% and 0.120–0.274% of body weight respectively across related species, indicating optimized cross-sectional areas for balanced force application to the pygostyle during locomotion.^[43] This setup ensures even load bearing, preventing localized stress concentrations observable in histological sections of attached tendons.^[4]

Role in Aerodynamics and Locomotion

The pygostyle anchors the rectrices, enabling birds to fan and adjust the tail as a dynamic control surface that modulates lift, drag, and stability during flight. By providing a rigid, low-mass terminus to the vertebral column, it allows precise manipulation of tail conformation to counteract perturbations, generating corrective moments for pitch and yaw, especially critical in low-speed phases like takeoff and landing where wing-generated control is limited.^[1] Wind-tunnel experiments on avian-inspired tail models quantify this function, showing that deployed tail feathers increase drag coefficients by up to 20-30% at angles of attack typical of perching or ascent, while asymmetric fanning produces yawing moments proportional to sideslip angle, enhancing directional stability without substantial mass penalty. Kinematic studies of live birds, such as pigeons and hummingbirds, further confirm that pygostyle-supported tail movements contribute 10-15% of total aerodynamic torque in maneuvering flight, minimizing energy expenditure by leveraging the tail's position posterior to the center of gravity.^[44]^[45]^[46] In flightless ratites like ostriches and emus, the pygostyle persists but exhibits reduced size and feather integration relative to volant birds, shifting its primary contribution from aerial control to terrestrial balance during bipedal locomotion, where tail undulation aids in counteracting lateral sway at speeds exceeding 50 km/h. This contrast underscores the pygostyle's adaptive optimization for flight in most taxa, with fusion enabling a compact structure that prioritizes control surface efficacy over extensibility, as deviations in caudal morphology correlate with flight capability across avian clades.^[47]^[4]

Paleontological and Scientific Debates

Fossil Evidence and Examples

The earliest documented occurrence of a pygostyle in the avian lineage is in Baminornis zhenghensis, a basal bird from the Late Jurassic Shangshaximiao Formation in Sichuan Province, China, dated to approximately 160–145 million years ago (mya), representing a fusion of the distal caudal vertebrae into a compact structure supporting tail feathers.^[48] This specimen pushes the fossil record of pygostyles back by about 18 million years compared to prior Early Cretaceous examples, with micro-computed tomography (micro-CT) scans confirming the bone's integrity and distinguishing it from unfused precursors in more basal theropods.^[49] In non-avian theropods, pygostyle-like fusions appear sporadically and independently, such as in the therizinosauroid Beipiaosaurus inexpectus from the Early Cretaceous Yixian Formation (~125 mya), Liaoning Province, China, where the terminal caudal vertebrae exhibit partial co-ossification without associated rectricial feathers, suggesting convergence rather than direct ancestry to avian forms.^[27] Scansoriopterygid theropods, like Epidexipteryx and related taxa from the Late Jurassic Daohugou Beds (~160 mya), Inner Mongolia, display shortened tails with distally clustered but unfused vertebrae forming a pygostyle-like rod, revealed through detailed skeletal preparations, though lacking the full fusion seen in crown-group birds.^[50] Early Cretaceous avialans from the Jehol Biota (~130–120 mya) provide transitional examples, including Jeholornis prima with a elongate tail of ~20 free caudal vertebrae terminating in a proximal fan of feathers but lacking a pygostyle, indicating an intermediate stage before full tail reduction.^[51] In contrast, pygostylian birds like Confuciusornis sanctus from the same Yixian Formation exhibit a rod-shaped pygostyle formed by 5–6 fused vertebrae, supporting a distal feather fan, as confirmed in multiple holotype specimens via radiographic imaging. Enantiornithine birds, such as Cruralispennia scissurrupium (~120 mya, Chaomidianzi Formation), display a ploughshare-shaped pygostyle convergent with ornithuromorphs, with micro-CT analyses of juvenile specimens revealing post-hatching fusion sequences akin to modern birds.^[14]^[3] Later Cretaceous fossils, including ornithuromorphs like Yixianornis grabaui from the Yixian Formation (~125 mya), show more derived, triangular pygostyles with expanded haemal spines for muscle attachment, facilitating enhanced tail control in flight, as detailed in museum-held specimens subjected to high-resolution scanning.^[29] These examples, spanning ~160–100 mya, underscore a mosaic pattern of pygostyle evolution, with independent fusions in multiple lineages documented through direct fossil preservation and advanced imaging techniques.^[2]

Homology, Convergence, and Evolutionary Implications

The pygostyle exhibits evidence of both homology within Avialae and convergent evolution across broader Maniraptora, complicating its role as a defining synapomorphy for Pygostylians. In early birds such as those from the Early Cretaceous Jehol Biota, the pygostyle is interpreted as a homologous structure supporting the reduction of caudal vertebrae to facilitate tail fan formation and aerodynamic control, distinguishing Pygostylians (including Enantiornithes and Euornithes) from long-tailed basal avialans like Archaeopteryx.^[52] However, fossil evidence from non-avialan theropods reveals independent origins of pygostyle-like fusion, as in Beipiaosaurus (a basal therizinosauroid from the Lower Cretaceous Yixian Formation), where distally co-ossified caudal vertebrae resemble the avian condition but likely served herbivorous balance functions rather than flight. This homoplasy is further supported by analogous structures in oviraptorosaurs and scansoriopterygids, indicating multiple evolutionary acquisitions of caudal fusion driven by selection for lightweight tails, not exclusively aerial locomotion.^[8] Within Enantiornithes, pygostyle morphology shows convergence on ornithuromorph-like plough-shaped forms, as in Cruralispennia (Early Cretaceous), where proximal forking and ventral processes evolved independently to anchor rectrices, despite differing phylogenetic positions from crown-group birds.^[14] Such instances challenge strict homology across Pygostylians, as enantiornithine pygostyles often retain elongate, forked proximal ends unlike the compact, triangular form in Ornithothoraces, suggesting adaptive convergence tied to fan-shaped tail feathers rather than shared ancestry alone.^[53] Empirical morphometric analyses of caudal skeletons further demonstrate shape convergence in diving neornithines, where pygostyle robustness correlates with underwater propulsion demands, independent of pygostyle origins in aerial clades.^[11] These patterns imply a polyphyletic or homoplastic evolutionary history for the pygostyle, undermining assumptions of gradual, unidirectional progression from theropod tails to avian sterna in bird-dinosaur transitions. Presence in non-volant therizinosaurs critiques over-reliance on pygostyle-feather co-evolution as diagnostic of powered flight, as fusion may primarily reflect mass reduction pressures in diverse ecologies, with aerial refinements as secondary adaptations in avialans.^[2] Fossil gradients from unfused theropod caudals to fused avian forms support mosaic evolution over strict gradualism, with 2017–2024 studies emphasizing functional causality—e.g., skeletal support for rectricial bulbs—over phylogenetic determinism, as abrupt appearances in basal records (e.g., Late Jurassic Baminornis with pygostyle) highlight rapid selective sweeps rather than incremental steps.^[8] ^[48] This convergence underscores causal realism in tail evolution, prioritizing biomechanical constraints like center-of-mass shifts for flyers over assumed linear homology from non-avian dinosaurs.

Etymology and Cultural Aspects

Terminological Origins

The term pygostyle originates from the Ancient Greek roots pygē (πυγή), denoting "rump" or "buttocks," combined with stylos (στῦλος), meaning "pillar" or "column," yielding a literal translation of "rump pillar" to describe the fused, supportive structure at the bird's tail base.^[54]^[55] This nomenclature emerged in the mid-19th century amid advancements in comparative anatomy, with the earliest English attestations recorded between 1870 and 1875, reflecting efforts to standardize avian skeletal terminology through Greco-Latin derivations for precision in scientific discourse.^[54] Prior to widespread adoption of pygostyle, early ornithological descriptions often employed more rudimentary or regionally variable terms, such as references to the "fused caudal vertebrae" or descriptive phrases emphasizing its terminal, pillar-like form in bird skeletons, as seen in 18th- and early 19th-century anatomical treatises.^[56] Alternative designations like "ploughshare bone," alluding to its flattened, blade-like shape in some species, appeared sporadically in older texts to evoke its supportive role akin to a plow's cutting edge, though this usage remained informal and less precise than the etymologically grounded pygostyle.^[8] Similarly, "uropygial bone" occasionally surfaced in contexts linking it to the uropygium (the rump region housing the preen gland), highlighting its positional integration rather than distinct morphology, but such terms yielded to pygostyle as phylogenetic systematics emphasized homologous structures across Aves.^[57] The shift toward pygostyle as the dominant term paralleled broader trends in 19th-century zoology, where nomenclature transitioned from ad hoc descriptive labels—often tied to gross morphology or function—to systematic compounds facilitating cross-species comparison and evolutionary analysis, without implying unsubstantiated homologies in non-avian theropods.^[55] This evolution underscored a commitment to terminological consistency in ornithology, enabling clearer documentation of skeletal reductions in avian tails post-Archaeopteryx.^[2]

Culinary and Historical Uses

The pygostyle, known colloquially as the "pope's nose," "parson's nose," or "bishop's nose," refers to the fatty, triangular nub at the base of the tail in poultry such as chickens, turkeys, and ducks, consisting of the fused caudal vertebrae surrounded by adipose tissue and skin.^[58] In culinary preparations, it is typically consumed as part of roasted or whole birds, prized by some for its crispy skin and rich flavor after cooking, though often left intact during carving due to its small size and high fat content.^[59] Nutritional analysis of similar poultry tail regions indicates elevated levels of fat and collagen; for instance, a comparable serving of turkey tail provides approximately 13 grams of protein and 40 milligrams of calcium, while chicken skin in the area contributes significant collagen alongside 20 grams of fat per 50-gram portion.^[60]^[61] Historically, the term "parson's nose" emerged in British English by the late 18th century, reflecting cultural associations with clerical figures, and the part has been eaten across European and Anglo-American traditions without documented widespread taboos, appearing in household roasts and anecdotal family preferences.^[62] In other cultures, such as Japanese yakitori, the equivalent "bonjiri" (fatty tail base) is grilled as a skewered delicacy for its juicy texture.^[63] Variations like "sultan's nose" suggest broader Eurasian naming, but consumption patterns emphasize its role as an incidental, flavorful byproduct rather than a standalone dish in most Western contexts. In modern poultry processing, the pygostyle region forms part of the offal or carcass remnants, often separated during evisceration and either rendered for fats, incorporated into pet foods, or discarded, with byproduct yields from broiler processing averaging 10-15% of live weight across categories like heads, feet, and viscera, though tail-specific data remains limited to general waste streams.^[64]^[65] Empirical studies on utilization highlight its potential in value-added products like hydrolyzed proteins, but routine culinary retention occurs primarily in whole-bird markets rather than industrial deboning lines.^[66]

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Aug 6, 2025 · ... fusion of the posteriormost caudal vertebrae. The thorough description in this study results in the emendation of diagnosis including the ...
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Oviraptorosaur Tail Forms and Functions - BioOne
Jan 4, 2013 · The tail is complete, composed of 33 vertebrae. The terminal three (or possibly two) caudal vertebrae are fused into a pygostyle (Fig. 4).
[29]
A new clade of basal Early Cretaceous pygostylian birds ... - PNAS
Sep 24, 2018 · We report the second most basal clade of the short-tailed birds (Pygostylia) from the Early Cretaceous. The new family Jinguofortisidae exhibits a mosaic ...
[30]
A bizarre Early Cretaceous enantiornithine bird with unique crural ...
Jan 31, 2017 · ... dorsal and ventral margins of the pygostyle are smooth and unbroken (Fig. ... The neck feathers are longer along the dorsal margin than the ...
[31]
Anatomy of the primitive bird Sapeornis chaoyangensis from the ...
Mar 2, 2017 · Sapeornis has, among many more derived features, a pygostyle, a furcula with a hypocleidum, a better developed humeral head and deltoid crest, a ...
[32]
A new confuciusornithid bird with a secondary epiphyseal ... - Nature
Dec 21, 2022 · The confuciusornithids are the earliest known beaked birds, and constitute the only species-rich clade of Early Cretaceous pygostylian birds
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Coevolution of caudal skeleton and tail feathers in birds - PubMed
Aug 20, 2014 · The structure of the tail feathers and the pygostyle each evolve in response to functional demands of differing locomotor behaviors. Here, I ...
[34]
Coevolution of Caudal Skeleton and Tail Feathers in Birds
Aug 20, 2014 · port the hypothesis of coevolution of caudal feathers and the pygostyle. Birds with different tail fan shapes (forked, graduated, or square).
[35]
An Early Cretaceous enantiornithine bird with a pintail - ScienceDirect
Nov 8, 2021 · The contrasting tail plumage between Early Cretaceous Jehol enantiornithines and ornithuromorphs represents a manifestation of the complexity ...
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New enantiornithine bird with an aerodynamic tail found in China
Jan 4, 2016 · However, similarities between the pygostyle in pengornithid enantiornithines, ornithuromorphs, and Sapeornis – which also has an aerodynamic ...
[37]
Morphological coevolution of the pygostyle and tail feathers in Early ...
Dec 10, 2023 · 摘要: The transformation from a long reptilian tail to a shortened tail ending in a pygostyle and accompanied by aerodynamic fanning rectrices ...
[38]
Distribution of Motoneurons Innervating Tail Muscles in the Pigeon
The movements of rectrices and pygostyle in pigeon are controlled mainly by following 6 muscles: M. levator caudae (MLVCD), M. depressor caudae (MDCA), M. ...
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[PDF] Morphological Aspects of Avian Tail Movements
the tail if both muscles act unilaterally. M. depressor caudae.--It is the deepest muscle that lies on the ventral surface of the free caudal.
[40]
An Enantiornithine with a Fan-Shaped Tail, and the ... - PubMed
In extant birds, this is typically associated with a fleshy structure called the rectricial bulb that secures the tail feathers (rectrices) [1]. The bulbi ...
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Bird Structure and Function - Advanced | CK-12 Foundation
The final few vertebrae are fused into a structure called a pygostyle. The pygostyle supports the tail feathers and muscles. The ribs are flattened, and the ...Missing: definition | Show results with:definition
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[PDF] ASC-202: Avian Skeletal System - Extension Publications
The tail has been reduced to a very short section of fused bones called the pygostyle ... The bones of birds are also lighter in weight than those of their ...<|separator|>
[43]
Tail myology and flight behaviour: Differences between caracaras ...
Feb 25, 2019 · In all species, the muscles depressor caudae and levator caudae had the highest mass values (0.028%–0.329% and 0.120%–0.274%, respectively) ...
[44]
Tail effects on yaw stability in birds - ScienceDirect.com
It is the purpose of this paper to show that tails of birds can generate yawing moments in case of a sideslip disturbance and, thus, contribute to stability in ...Missing: fanning | Show results with:fanning
[45]
Sharp turning maneuvers with avian-inspired wing and tail morphing
Nov 24, 2022 · We experiment in wind tunnel and flight tests how such morphing capabilities increase the roll rate and decrease the turn radius - two measures ...Avian-Inspired Drone · Morphing Drone · Roll And Yaw Moment Wind...
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(PDF) The Experimental Study of Tail Shape for Hummingbird-like ...
The results show that the lift generated by the tail is independent of the fan angle in this study, whereas pitch moment is related to the fan angle. Also, the ...
[47]
[PDF] Flying vs flightless birds
Fused caudal vertebrae. • Serve as a docking platform the retricial bulb and associated muscles and feathers for controlling tail fan and contour shape.
[48]
New fossils imply a deeper origin of modern birds in the Mesozoic
Jun 9, 2025 · Baminornis zhenghensis is now the earliest known bird exhibiting a pygostyle, pushing back the fossil record of pygostyles by approximately 18 ...
[49]
China Jurassic fossil discovery sheds light on bird origin - EurekAlert!
Feb 12, 2025 · Notably, Baminornis zhenghensis has a short tail ending in a compound bone called the pygostyle, a feature also observed in living birds. " ...
[50]
[PDF] Theropoda)Zhongornis haoaeas a scansoriopterygid-like non-avi
A similar structure is present in Epidexipteryx, Similicaudipteryx, and several Late ... vertebrae form a pygostyle-like structure but remain unfused (Zhang et al ...Missing: Scansoriopteryx | Show results with:Scansoriopteryx
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Unique caudal plumage of Jeholornis and complex tail evolution in ...
Oct 7, 2013 · Jeholornis is unique among all known birds in possessing a proximally positioned tail fan that is not associated with a pygostyle. Gatesy ...
[52]
The Origin and Diversification of Birds - ScienceDirect.com
Oct 5, 2015 · Early birds diversified throughout the Jurassic and Cretaceous, becoming capable fliers with supercharged growth rates, but were decimated at ...
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An Enantiornithine with a Fan-Shaped Tail, and the ... - Cell Press
Dec 31, 2015 · In extant birds, this is typically associated with a fleshy structure called the rectricial bulb that secures the tail feathers. (rectrices) ...
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PYGOSTYLE definition in American English - Collins Dictionary
Word origin. [1870–75; ‹ Gk pȳgo- (comb. form repr. pȳge᷄ rump) + stŷlos pillar]This word is first recorded in the period 1870–75. Other words that entered ...Missing: etymology | Show results with:etymology
[55]
pygostyle, n. meanings, etymology and more
pygostyle is a borrowing from Greek, combined with an English element. Etymons: pygo- comb. form, Greek στῦλος. See etymology. Nearby entries. pygo- ...
[56]
Pygostyle - The Collaborative International Dictionary of English
Py go*style, n. [Gr. pygh the rump + ? a pillar.] (Anat.) The plate of bone which forms the posterior end of the vertebral column in most birds; ...
[57]
The Preen Gland - Ornithology
Dec 11, 2022 · Uropygial comes from Greek words meaning “rump” and pygostyle from the Greek meaning “rump pillar. ... Bird : Ornithological History Through Forty ...Missing: terminology | Show results with:terminology
[58]
Word Of The Day: Parson's Nose - Food Republic
Jun 8, 2015 · A parson's nose is the triangular stub where tailfeathers grow on poultry. ... Chances are if you're cooking, you'll be trying the pope's nose ( ...Missing: culinary | Show results with:culinary
[59]
This Is Why We Call A Turkey's Tailbone The Pope's Nose - Mashed
Jan 18, 2021 · Some favor the term pope's nose to describe a turkey's tailbone while others call it a parson's nose instead. Why does a bird's butt-bone ...Missing: culinary | Show results with:culinary
[60]
Chicken butt is such a delicacy it has fancy names | campus.sg
Mar 29, 2021 · ... chicken breast, and a serving of turkey tail provides 13g of protein and 40mg of calcium. Pope's Nose sold in Samoa. Chinese: 七里香 (7 Mile ...
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Utilization of byproducts and waste materials from meat, poultry and ...
Available information pertaining to the utilization of by-products and waste materials from meat, poultry and fish and their processing industries has been ...
[65]
[PDF] Processing of poultry byproducts and their utilization in feeds
Processed byproduct yields and potential supply from principal types of poultry. [Live-weight basis, 1958production]. Poultry type. Processed byproduct yields ( ...
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Thermal hydrolysis of poultry byproducts for the production of ... - NIH
Feb 19, 2025 · The objective of this research is to test the effect of processing DAF solids obtained directly from the DAF unit versus course-filtered DAF ...