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Paraves

Paraves is a clade of maniraptoran theropod dinosaurs within the larger group Coelurosauria, phylogenetically defined as the stem-based taxon comprising all coelurosaurs more closely related to Passer domesticus (the house sparrow) than to Oviraptor philoceratops or Ornithomimus edmontonicus ^[1]. This clade encompasses the immediate relatives of birds and modern Aves, including the subclades Deinonychosauria (which contains Dromaeosauridae and Troodontidae) and Avialae, marking a critical evolutionary branch in the transition from non-avian dinosaurs to birds. Notable taxa include Deinonychus antirrhopus, Velociraptor mongoliensis, Troodon formosus, Microraptor zhaoianus, Sinornithosaurus millenii, and Archaeopteryx lithographica, alongside over 11,000 extant bird species (as of 2025) ^[2]. Paraves originated in the Late Jurassic and persisted for over 160 million years into the present, with fossils primarily from Cretaceous formations such as the Djadokhta in Mongolia and the Jehol Group in China ^[3]. Key synapomorphies defining the clade include pennaceous feathers on the body and limbs, fused parietals, an L-shaped scapulocoracoid, a symmetrical furcula, and modifications for enhanced forelimb mobility, such as a humerus longer than the scapula ^[1]. These features reflect adaptations toward powered flight, with basal paravians like Microraptor exhibiting "four-winged" gliding capabilities via elongated feathers on all limbs, while avialans show further refinements like asymmetrical vanes and reduced tails. The evolutionary significance of Paraves lies in its role as the nexus for avian origins, bridging non-avian theropods and modern birds through innovations in integument, skeletal pneumatization, and body size reduction—spanning four orders of magnitude in mass over approximately 15 million years. Phylogenetic analyses, incorporating up to 111 taxa and 474 morphological characters, consistently recover Paraves as a monophyletic group sister to oviraptorosaurs or therizinosauroids within Maniraptora, supported by at least 12 unambiguous synapomorphies ^[1]. Numerous non-avian paravian species are known, providing a rich fossil record that illuminates feather evolution, predatory behaviors, and the ecological radiation of feathered dinosaurs across Asia, North America, and Europe ^[3].

Anatomy

Skeletal structure

Paraves is characterized by several key skeletal features distinguishing it from more basal maniraptorans, including a large promaxillary fenestra, a T-shaped lacrimal, a lateral posteriorly expanding longitudinal groove on the dentary, manual phalanx IV-2 shorter than IV-1, a short ischium with distally located obturator process and posterodistal process, and an extensible pedal phalanx II-2.^[3] These traits support the monophyly of the clade encompassing deinonychosaurians and avialans. A recent study on the cranial anatomy of Anchiornis huxleyi provides additional details on skull evolution, highlighting features transitional to avian conditions.^[4] The furcula, or wishbone, in Paraves exhibits a distinctive boomerang-shaped morphology, often U-shaped with short, broad, and dorsoventrally expanded clavicular rami that are nearly symmetrical and sometimes pneumatic, as observed in taxa like Buitreraptor (MPCA 245) and Velociraptor mongoliensis.^[1]^[5] This structure articulates medially with the scapulocoracoid complex and serves to stabilize the shoulder girdle by providing robust attachment sites for flight-related muscles, such as the m. pectoralis, thereby enhancing forelimb excursion during locomotion or aerial behaviors.^[5]^[6] Cranial features of Paraves include a flexible quadrate with a dorsally displaced triangular process, enabling prokinetic movement, and a large antorbital fenestra that is vertically oriented with a broadly rounded anterior border formed by the lacrimal and nasal.^[1] These adaptations facilitate kinetic skull movement by allowing independent flexion of the rostrum relative to the braincase, a trait that originated basally within Paraves and supported enhanced feeding efficiency through improved jaw mobility and reduced skull mass.^[7]^[8] In the axial skeleton, Paraves display heterodont dentition with serrated teeth of varying sizes, such as larger first and second premaxillary teeth compared to posterior ones, exemplified by Deinonychus antirrhopus (YPM 2505) with 15 maxillary, 4 premaxillary, and 16 dentary teeth where posterior denticles are nearly twice as large as anterior ones.^[1] The cervical vertebrae typically number 10–11, providing elongated and flexible necks for greater maneuverability, as seen across taxa like Sinovenator and Mei.^[1]^[9] A derived trait within avialans, the pygostyle, involves fusion of the distal caudal vertebrae into a single structure, as documented in basal forms like Confuciusornis sanctus (IVPP V11374), which anchors tail feathers and marks an adaptation for aerodynamic control.^[10]

Forelimbs and wings

The forelimbs of paravians exhibit significant elongation relative to the hindlimbs, a key adaptation for aerial locomotion, with the manual digits particularly extended to support feather anchorage. In basal paravians such as Anchiornis and Microraptor, digits II and III are notably lengthened, forming the primary structural base for attaching flight feathers, while digit I remains shorter and more claw-like.^[9]^[11] This configuration allows for a broadened wing surface, with primary remiges anchoring primarily along digits II and III, enhancing stability during gliding or flapping.^[5] Pennaceous feathers covering the paravian arms form asymmetrical vanes critical for generating lift, consisting of a central rachis from which barbs extend, interconnected by barbules with hooklets that create a cohesive, airfoil-like structure. In paravians, these feathers evolved asymmetrical vanes at the clade's base, with the leading-edge vane narrower than the trailing edge to optimize airflow and reduce drag during aerial maneuvers.^[12]^[13] The rachis provides rigidity, while the barbule arrangements ensure vane integrity under aerodynamic stress, as seen in the densely packed barbs of arm feathers that form broad, planar surfaces.^[14] The shoulder girdle in paravians features specialized adaptations for expansive forelimb motion, including a strut-like coracoid that articulates firmly with the scapula to transmit forces efficiently during wing beats. The scapula is elongated, often forming an L-shaped complex with the coracoid, which positions the glenoid fossa laterally and subvertically, permitting a wide range of anteroposterior and elevational movements essential for powered or gliding flight.^[5]^[15] These features, observed in taxa like Microraptor and Archaeopteryx, support enhanced mobility compared to more basal theropods.^[5] Wing loading in paravians varies, with Archaeopteryx estimated to have relatively low values enabling potential powered flight, in contrast to non-avialan paravians like Microraptor suited more for gliding. This reflects differences in wing proportions and body size across the clade. Fossil evidence from Microraptor exemplifies these traits, with exceptionally preserved feather imprints revealing four-winged configurations where forelimb primaries (~10 asymmetrical remiges) and secondaries (~17–21 symmetrical feathers) form layered, V-shaped wings for gliding, supported by soft tissues like the propatagium.^[16] These imprints show close-vaned structures curving medioposteriorly, indicating aerodynamic refinement for controlled descent.^[16]

Hindlimbs and pedal features

The hindlimbs of paravians exhibit specialized adaptations for agile terrestrial locomotion and predation, with the pes (foot) displaying a subarctometatarsalian or fully arctometatarsalian condition in which metatarsal III is pinched proximally between metatarsals II and IV, reducing the overall foot width while enhancing rigidity and cursorial efficiency. This configuration, observed in troodontids and dromaeosaurids, allows for a more streamlined stride during rapid movement, with metatarsal III often comprising a significant portion of hindlimb length relative to the femur and tibia.^[17]^[18] In microraptorines and unenlagiines, the condition is subarctometatarsalian, providing transitional flexibility between broader theropod feet and the more compact avian tarsometatarsus. Recent studies highlight diversity in hindlimb feathering among early paravians, including asymmetrical vanes on leg feathers in Microraptor, contributing to four-winged gliding.^[19]^[1] A hallmark feature of deinonychosaurian paravians is the enlarged, recurved ungual on pedal digit II, forming the iconic sickle claw that is hypertrophied relative to other pedal unguals and held retracted off the ground during locomotion via a hyperextensible metatarsophalangeal joint. This claw, strongly curved and falciform, measures several centimeters in length in taxa like Deinonychus and Velociraptor, enabling powerful grasping during predatory strikes. Fossil evidence indicates the presence of a keratinous sheath extending the functional length and sharpness of the ungual, as preserved organic material on pedal phalanges in specimens such as Rahonavis ostromi and basal paravians shows immunoreactivity consistent with beta-keratin composition.^[20]^[21]^[22] The fibula in paravians is notably reduced, adopting a splint-like morphology that tapers distally and attaches closely to the tibia, contributing to enhanced leg stability by buttressing the tibia's lateral surface without bearing significant weight. In dromaeosaurids such as Rahonavis and Unenlagia, the fibula's midshaft diameter is less than one-fifth that of the tibia, and it lacks extensive contact with the calcaneum, reflecting lightweight adaptations for speed. This reduction, combined with proximal fusion of the astragalus and calcaneum to the tibia in many paravians, optimizes force transmission during bipedal strides.^[1]^[23] The pubic boot, a distal expansion of the pubis forming a sub-triangular or spatulate structure, is present in several paravian clades including unenlagiids, microraptorines, and anchiornithines, potentially providing anchorage for abdominal musculature involved in posture and locomotion. In these taxa, the boot projects posteriorly and supports the retroverted pubis characteristic of maniraptorans, though its role in reproductive efficiency remains speculative without direct evidence of associated soft tissues.^[3] Fossil specimens of Deinonychus antirrhopus, such as those from the Cloverly Formation, reveal a hyperextensible ankle joint facilitated by the mesotarsal articulation between the tibia and proximal tarsals, allowing extreme dorsiflexion of the foot during raptorial maneuvers. Musculoskeletal modeling of the Deinonychus hindlimb demonstrates that this configuration, combined with the arctometatarsal pes, maximizes leverage for slashing or pinning strikes, with the ankle's flexibility enabling rapid deployment of the sickle claw.^[24]

Size and body plan variation

Paraves exhibit a broad range of body sizes, from diminutive forms like Anchiornis huxleyi, which measured approximately 0.5 meters in length and weighed under 1 kg, to larger dromaeosaurids such as Utahraptor ostrommaysorum, reaching up to 7 meters in length and around 500 kg.^[25]^[26]^[27] Body plan proportions vary significantly across the clade, particularly in tail morphology, where non-avialan paravians typically possess long, flexible tails comprising over half the total body length for balance and maneuverability, in contrast to the shortened, stiffened pygostylian tails of avialans that support feathered fans for aerodynamic control.^[28]^[29] Allometric scaling influences physiological traits, with smaller paravians displaying bone histology indicative of higher metabolic rates, such as dense vascularization and rapid deposition rates in cortical bone, compared to larger relatives with slower growth lines suggesting more moderate metabolism.^[30] Troodontids exemplify encephalization variation, possessing brains with an encephalization quotient approximately five times greater than that of other theropods, reflecting expanded cerebral regions relative to body size despite their generally small stature.^[31]^[32] Evidence of intraspecific body plan variation comes from growth series in fossils of Sinornithosaurus, where multiple specimens reveal ontogenetic shifts in limb proportions and feather development, indicating flexibility in size and morphology during individual development.^[33]^[34]

Classification

Taxonomic history

The taxonomic history of Paraves reflects a progression from fragmented understandings of theropod dinosaurs to a cohesive phylogenetic framework, driven by key fossil discoveries and cladistic analyses. In the mid-20th century, non-avialan theropods like dromaeosaurids were often viewed as primitive or distantly related to birds, with limited resolution in their placement among coelurosaurs. This changed with John H. Ostrom's 1969 description of Deinonychus antirrhopus from the Early Cretaceous of Montana, where he recognized its unusual features—such as the enlarged sickle-shaped claw and bird-like skeletal proportions—and classified it within Coelurosauria, elevating awareness of maniraptoran diversity.^[35] Ostrom's subsequent work in the 1970s further bridged birds and theropods. His 1976 analysis of Archaeopteryx emphasized osteological similarities to Deinonychus, including the furcula, pelvic structure, and forelimb morphology, proposing that Archaeopteryx represented an early avian derivative of small coelurosaurian dinosaurs closely allied with dromaeosaurs.^[36] This revived the dinosaur-bird hypothesis, challenging earlier views that separated birds from reptiles. The formal establishment of Paraves came in 1998, when Paul C. Sereno defined it as a stem-based clade encompassing all maniraptorans more closely related to Neornithes than to Oviraptor philoceratops, thereby uniting Avialae (bird-like theropods including Archaeopteryx), Dromaeosauridae, and Troodontidae on the basis of shared traits like reduced manual digits and enhanced arm mobility.^[37] Prior nomenclature, such as Jacques Gauthier's 1986 stem-based definition of Avialae (all dinosaurs closer to birds than to Deinonychus), had partially overlapped with this concept but focused more narrowly on ornithurine descendants; Sereno's Paraves provided a broader, explicit framework for the avian radiation. In a 2001 symposium honoring Ostrom, Gauthier and Kevin de Queiroz refined avian terminology, redefining Avialae to include all dinosaurs with feathered wings adapted for flapping flight and their descendants, while distinguishing it from the more inclusive Paraves to avoid conflating flight origins with deeper theropod affinities.^[38] The early 2000s brought transformative fossil evidence from China, led by Xu Xing, that solidified Paraves' monophyly. Discoveries of feathered non-avialan theropods, such as the four-limbed glider Microraptor zhaoianus in 2000 and additional "four-winged" dromaeosaurids in 2003, revealed pennaceous feathers, aerodynamic adaptations, and skeletal features aligning closely with avialans, confirming the clade's shared integumentary and locomotor traits beyond Europe and North America. Ongoing debates centered on enigmatic taxa like Scansoriopterygidae, known from Late Jurassic specimens such as Epidexipteryx (described in 2008). Initially placed outside Maniraptora due to their elongated third finger and membranous wing structures, phylogenetic analyses in the 2010s and beyond have debated their position, with some recovering them within Paraves as basal avialans based on shared paravian synapomorphies like furcula presence and feather impressions, while others suggest affinities outside the clade or with oviraptorosaurs; expanded datasets incorporating Chinese fossils continue to show instability as of 2025.^[3]

Phylogenetic position

Paraves represents a derived clade within the maniraptoran theropods, positioned as the sister group to Oviraptorosauria in the broader framework of Maniraptora, a subdivision of Coelurosauria. This relationship is consistently recovered in large-scale cladistic analyses incorporating hundreds of morphological characters across theropod taxa, emphasizing Paraves' role in the transition toward avian dinosaurs. The clade is defined as a stem-based taxon including all maniraptorans closer to Neornithes than to Oviraptor philoceratops (Sereno, 1998), though some studies use a node-based definition as the most inclusive group containing Archaeopteryx, Dromaeosaurus, and Troodon excluding oviraptorosaurs.^[39]^[40]^[37] Phylogenetic trees derived from parsimony-based analyses depict Paraves diverging into two primary lineages: Deinonychosauria, which encompasses Dromaeosauridae (e.g., Velociraptor, Microraptor) and Troodontidae (e.g., Mei, Zanabazar), and Avialae, which includes basal forms such as Jeholornis alongside more crownward groups like Euornithes and Ornithurae (modern birds and their close relatives).^[39]^[40] Diagnostic synapomorphies uniting Paraves include the enlargement and robustification of the alular digit (manus digit I), which supports enhanced manual dexterity and potential aerodynamic function; the development of uncinate processes on the dorsal ribs, aiding in thoracic stabilization during locomotion or flight; and the presence of a secondary antorbital fenestra, a cranial opening associated with lightweight skull construction. Additional shared features encompass a retroverted pubis in derived members, contributing to a more avian-like pelvic girdle, and large, posteriorly projecting axial epipophyses on the vertebrae for enhanced neck mobility. These characters are scored in matrices that resolve Paraves with strong clade support, such as Bremer indices of 3 and jackknife values indicating low collapse potential in reduced consensus trees.^[39] Quantitative support for Paraves' monophyly and internal topology is robust in seminal datasets; for instance, Brusatte et al. (2015) report bootstrap values of 80–90% for the Paraves node and its major subclades in time-calibrated analyses of theropod evolution, reflecting consistent recovery across 230 taxa and 457 characters despite variations in character weighting. Updates in subsequent studies, including those incorporating new paravian fossils up to 2025, maintain this topology with similar high support (e.g., >70% bootstrap for Deinonychosauria).^[40] Controversies persist regarding the exact placement of certain paravian taxa, such as Pyroraptor olympius, which fragmentary remains have led some analyses to position as a basal dromaeosaurid within Deinonychosauria, while others suggest closer avialan affinities due to inferred flight-related traits; however, most comprehensive matrices favor its inclusion in Eudromaeosauria with moderate support (Bremer index ~2).^[39]

Included taxa

Paraves encompasses several major subgroups, each characterized by distinct morphological features that reflect their adaptations and evolutionary roles within the clade. The primary divisions include Avialae, Deinonychosauria, and various basal paravians.^[41] Avialae comprises Archaeopteryx and all modern birds (Aves), along with numerous extinct stem-group members such as enantiornithines from the Cretaceous. This clade is defined by key synapomorphies including the presence of a pygostyle—a fused terminal caudal vertebra that supports tail feathers—and a strut-like coracoid bone that enhances shoulder girdle stability for flight-related functions. Its diversity spans from Jurassic stem-birds like Archaeopteryx, known from Solnhofen limestone deposits, to highly diverse Cretaceous forms like enantiornithines, which dominated Mesozoic avifaunas with over 80 described species exhibiting varied body sizes and ecological niches.^[41]^[42] Deinonychosauria includes the families Dromaeosauridae and Troodontidae, representing non-avialan paravians specialized for predatory lifestyles. Dromaeosauridae, exemplified by Velociraptor mongoliensis from Late Cretaceous Mongolia, features cursorial adaptations such as elongated hindlimbs with a raptorial sickle claw on pedal digit II, enabling agile terrestrial hunting. Troodontidae, including Mei long from Early Cretaceous China, is distinguished by enhanced cranial flexibility, evidenced by kinetic skull elements like a mobile quadratojugal joint, which likely facilitated precise prey manipulation and supported their relatively large brains relative to body size.^[41] Basal paravians form a paraphyletic assemblage of early-diverging members outside the crown clades, highlighting the transitional diversity within Paraves. Anchiornithidae, such as Anchiornis huxleyi from Late Jurassic China, are recognized for their four-winged body plan with pennaceous feathers on both fore- and hindlimbs, suggesting gliding capabilities in arboreal or forested environments. Scansoriopterygidae, including Yi qi from Middle Jurassic China, exhibit arboreal climbing adaptations, notably an elongated third manual digit that may have supported patagial membranes for gliding or scanningorial locomotion; their exact position within or relative to Paraves remains debated.^[41] Approximately 50 non-avian paravian species have been described, underscoring the clade's extensive Mesozoic radiation, with approximately 70% originating from Late Jurassic to Early Cretaceous deposits in Asia, particularly the Yanliao and Jehol biotas of northeastern China. This Asian concentration reflects exceptional Lagerstätten preservation that has revealed feathered specimens and informed phylogenetic reconstructions.^[41] Groups like Oviraptorosauria, including oviraptorids, are excluded from Paraves based on phylogenetic analyses showing they lack core paravian synapomorphies such as the enlarged forelimbs with elongated manual digits and the hyperextensible pedal digit II, instead aligning with more basal maniraptorans in broader theropod trees.^[41]

Evolutionary history

Origins and early diversification

Paraves likely originated in the Middle Jurassic, with phylogenetic analyses indicating a divergence from other maniraptorans around 170 Ma, coinciding with an acceleration in body size miniaturization that characterized the early evolution of the clade.^[43] This miniaturization trend among maniraptorans, reaching body sizes under 1 kg in basal paravians, may have been driven by predation pressures or adaptations to forested environments, as evidenced by the warm, humid ecosystems of the Yanliao Biota where early paravians are preserved. The earliest potential fossil records of Paraves come from Anchiornis-like forms in the Tiaojishan Formation of northeastern China, dated to approximately 160 Ma, representing the oldest known specimens of the clade.^[3] Ghost lineages inferred from cladistic analyses extend the inferred stem age of Paraves to about 170 Ma, filling gaps in the fossil record prior to the Late Jurassic radiation of coelurosaurs following recovery from the end-Triassic extinction.^[43]^[44] Early diversification of Paraves occurred by the Late Jurassic, with taxa radiating into arboreal, aerial, and terrestrial niches, facilitated by anatomical innovations such as elongated forelimbs and feathered appendages. This radiation is exemplified by the presence of protofeather-like structures in contemporaneous non-paravian dinosaurs like Tianyulong from around 160 Ma, suggesting a broader coelurosaurian experimentation with integumentary coverings that supported niche expansion in forested habitats. Molecular clock estimates place the crown-group divergence of Paraves between 170 and 160 Ma, aligning with the post-Triassic coelurosaur explosion and initial Pangaean distribution before continental breakup restricted later biogeography to primarily Laurasian realms.^[43]^[44]

Fossil record overview

The fossil record of Paraves is primarily known from exceptional Lagerstätten that preserve delicate structures such as feathers and soft tissues, with the Solnhofen Limestone in Germany standing out as a key Late Jurassic (~150 Ma) site. This fine-grained limestone, formed in anoxic lagoonal environments, has yielded multiple specimens of Archaeopteryx, including the renowned Berlin specimen (Humboldt Museum HMN MB.AV.101), which features a nearly complete articulated skeleton with impressions of primary flight feathers and other integumentary details.^[45]^[3] Similarly, the Early Cretaceous Yixian Formation (~125 Ma) in Liaoning Province, China, represents another pivotal deposit of volcanic ash and lacustrine sediments that facilitated the preservation of feathered paravians, such as the dromaeosaurid Sinornithosaurus millenii, whose holotype (IVPP V12719) includes filamentous integument covering much of the body.^[46]^[3] Beyond these Asian and European hotspots, notable discoveries include the dromaeosaurid Deinonychus antirrhopus from the Cloverly Formation in Montana, USA, with specimen AMNH 3015—a partial but informative skeleton collected in the 1960s—providing early insights into paravian skeletal robustness despite lacking soft-tissue preservation.^[35] Other significant sites encompass the Tiaojishan Formation in China, which has produced over 200 specimens of the anchiornithine Anchiornis huxleyi with detailed plumage, and the Djadochta Formation in Mongolia, yielding troodontids and dromaeosaurids in eolian dune settings.^[3] Preservation in these deposits favors small-bodied, feathered taxa due to the fine-grained, low-energy sedimentary environments of lakes and lagoons, which inhibit decay and scavenging, leading to an overrepresentation of arboreal or volant forms while undersampling larger, terrestrial paravians.^[3] Discovery trends show a marked increase since the 1990s, with a majority of paravian specimens originating from Asia, particularly China, where lagerstätten like Yixian have contributed to the description of dozens of new species, fundamentally reshaping understandings of paravian diversity. However, significant gaps persist, including scant records from the Middle Jurassic—prior to the oldest known fossils—where paravians likely originated but direct fossils are rare, and the Southern Hemisphere, limited mostly to isolated unenlagiine finds in Patagonia, Argentina, such as Unenlagia comahuensis and the recently described Diuqin lechiguanae from the Santonian (~85 Ma).^[47]^[3]

Temporal distribution

Paraves first appeared in the fossil record during the Middle to Late Jurassic, spanning approximately 160–145 million years ago (Ma), with early basal forms such as Aurornis xui from the Tiaojishan Formation of northeastern China representing some of the oldest known members of the clade. This period is characterized by small, feathered paravians exhibiting transitional features between non-avialan theropods and more derived birds.^[3] In North America, the Late Jurassic Morrison Formation (~155–145 Ma) has preserved incomplete paravian fossils, including elements attributed to Hesperornithoides miessleri, indicating an early presence of the group in Laurasian continental deposits.^[48] Following this initial emergence, Paraves underwent significant diversification during the Early Cretaceous (~145–100 Ma), reaching a peak in abundance and morphological variety within the Jehol Biota of northeastern China, dated to approximately 130–120 Ma, where exceptionally preserved specimens reveal advanced feathering and flight-related adaptations in taxa like microraptorines and early avialans.^[49] The Late Cretaceous (~100–66 Ma) saw continued paravian presence, particularly among larger dromaeosaurids such as Dakotaraptor steini from the Maastrichtian Hell Creek Formation of South Dakota, USA (~68–66 Ma), highlighting the persistence of non-avialan forms until the end of the Mesozoic. Non-avialan paravians abruptly vanished at the Cretaceous-Paleogene (K-Pg) boundary (~66 Ma), an event linked to the Chicxulub asteroid impact and associated environmental disruptions, including habitat destruction and climate shifts. In contrast, avian paravians—stem-birds and early crown-group representatives—persisted through the K-Pg mass extinction, with fossil evidence showing their survival in reduced numbers immediately following the event, while crown-group birds underwent rapid post-extinction diversification during the Paleogene.^[50]

Paleobiology

Locomotion and flight capabilities

Paravians exhibited a spectrum of locomotor strategies, ranging from quadrupedal gliding in basal forms to cursorial running in dromaeosaurids and powered flapping in avialans, reflecting biomechanical adaptations for diverse environments. Basal paravians like Microraptor utilized a four-winged planform, with elongated feathered forelimbs and hindlimbs forming a biplane-like configuration that facilitated stable gliding. Computer simulations of Microraptor flight performance indicate this morphology was adapted for undulatory "phugoid" gliding between trees, allowing controlled descent with lateral body movements for maneuvering. Physical models tested in wind tunnels have estimated a lift-to-drag ratio of 4.1:1 for Microraptor gui, corresponding to glide angles averaging 13.7° and enabling efficient aerial travel over moderate distances.^[51]^[52] Dromaeosaurids demonstrated pronounced cursorial adaptations suited to terrestrial sprinting, characterized by elongated hindlimbs with tibia-to-femur length ratios of approximately 1.2–1.3, which enhanced stride efficiency and agility. These proportions, observed in taxa such as Deinonychus and Dakotaraptor, correlate with biomechanical correlates of superior running ability, including reduced body mass relative to leg length and a digitigrade posture. Limb scaling and trackway analyses infer maximum speeds up to 40 km/h for mid-sized dromaeosaurids like Deinonychus, allowing rapid pursuit across open terrains.^[53]^[54]^[55] The transition to powered flight in avialans involved enhanced shoulder girdle mobility, permitting greater wing excursion during the downstroke to generate propulsive force. In basal avialans, the glenoid fossa's orientation and furcula-scapula articulation allowed abduction and depression of the humerus, optimizing power output from the pectoralis muscle. Muscle attachment scars on the sternum and coracoid, such as those for the pectoralis major, indicate substantial leverage for depressing the wing, with reconstructed musculature suggesting this configuration could produce aerodynamic forces comparable to early flapping in modern birds.^[56]^[5] Locomotor strategies varied between arboreal and terrestrial niches within Paraves. Scansoriopteryx, an arboreal scansoriopterygid, possessed a short, stiffened tail that likely functioned as a prop for balance and support during trunk climbing, analogous to woodpecker tails, facilitating ascent along vertical surfaces with its elongated manual digits. In contrast, troodontids exhibited terrestrial adaptations, including elongated metatarsals forming an arctometatarsal pes that enhanced foot flexibility and spring-like energy storage for leaping and rapid maneuvers over ground. These long metatarsals, combined with a nimble hallux, supported bounding gaits in pursuit or evasion scenarios.^[57] Biomechanical models have elucidated flight performance in early avialans like Archaeopteryx. Wind tunnel experiments on physical wing models demonstrate lift coefficients approaching 1.2 during downstroke simulations, highlighting the role of asymmetric feather vanes and wing camber in generating sufficient lift for short bursts of powered flight or controlled glides from elevated starts. These analyses underscore how subtle morphological features, such as alula-like structures, mitigated stall risks and improved aerodynamic efficiency.^[58]

Diet and predatory adaptations

Members of Paraves, particularly deinonychosaurs, possessed ziphodont dentition adapted for slashing flesh, featuring laterally compressed, recurved crowns with fine serrations that facilitated efficient tearing of prey tissues. These serrations typically exhibited densities of 5-10 per millimeter, enhancing the cutting efficiency during predation. In avialans, dentition evolved toward more versatile forms, with some taxa developing broader crowns and grinding occlusal surfaces suited to processing a wider range of foods beyond strict carnivory.^[59]^[60]^[61] Predatory behaviors in paravians included cooperative strategies, as evidenced by bonebeds containing multiple Deinonychus individuals alongside disarticulated Tenontosaurus skeletons, suggesting group hunting or scavenging of large herbivores. The iconic sickle-shaped claw on pedal digit II of dromaeosaurids served primarily to pin and restrain struggling prey, with biomechanical analyses estimating it could exert forces up to approximately 500 N to secure victims during attacks. These adaptations complemented agile locomotion, enabling effective pursuit and capture.^[62]^[24] Cranial kinesis provided key predatory advantages, with flexible jaw mechanisms allowing expansive gapes of up to 80° to engulf larger or more agile prey items. In troodontids, prokinetic motion—where the upper jaw flexed relative to the braincase—further enhanced this capability, permitting rapid and precise strikes in low-light or confined environments.^[63] Dietary habits varied across Paraves, reflecting ecological diversification. Small avialans such as Jeholornis displayed omnivorous tendencies, with fossilized gut contents preserving intact seeds from fruits, indicating consumption of plant material alongside animal prey for seed dispersal. Piscivory characterized certain avialans, including Hesperornis, whose conical teeth and robust jaws were specialized for grasping and consuming fish in marine settings. Stable isotope analyses of paravian fossils, particularly δ¹³C values, reveal mixed terrestrial and aquatic dietary signals in some taxa, underscoring opportunistic foraging strategies that bridged habitats.^[64]^[65]^[66]

Reproduction and growth

Paraves laid elongated, hard-shelled eggs measuring approximately 10-15 cm in length, arranged in ovoid clutches of up to 24 eggs, with a calcite-based microstructure consisting of a mammillary inner layer and a squamatic outer layer similar to that of modern birds.^[67]^[68] Nesting evidence from troodontids includes clutches of over 20 eggs arranged in concentric rings, partially buried in substrate, suggesting brooding behavior with direct body contact for incubation.^[69] In oviraptorids, close relatives of paravians, multiple Citipati specimens preserve adults in a brooding posture with wings folded over the clutch and head tucked under a wing, consistent with thermoregulatory incubation; associated embryos exhibit advanced development, including curled postures indicative of late-stage hatching.^[70]^[71] Bone histology in paravians reveals rapid, avian-like growth characterized by fibrolamellar bone tissue, with von Bertalanffy growth constants (k) of 0.3-0.5 per year, enabling small taxa such as troodontids and dromaeosaurids to reach skeletal maturity in 1-2 years.^[72]^[73] Evidence for parental care in paravians includes incubation behaviors inferred from nest-associated troodontid specimens. Medullary bone, indicating calcium mobilization for eggshell formation in gravid females, has been identified in close relatives such as oviraptorids; however, brooding adults preserved on nests lack medullary bone, implying paternal incubation in some cases.^[74] Debates on hatchling development center on evidence for precociality, with troodontid nests showing no crushed eggshells or trampling traces, and well-ossified embryonic skeletons suggesting high mobility at hatching rather than altricial dependence.^[75]^[76]

References

[1]
A Review of Dromaeosaurid Systematics and Paravian Phylogeny
Aug 17, 2012 · Paraves. As coined and defined by Sereno (1997, then 1998 respectively), Paraves is a stem-based clade of derived coelurosaurs named to set ...
[2]
Comments on the Morphology of Basal Paravian Shoulder Girdle
May 24, 2021 · The present paper aims to review some important topics on pectoral girdle anatomy related to flight origins.
[3]
(PDF) Unusual pectoral apparatus in a predatory dinosaur resolves ...
Aug 6, 2025 · The furcula is a distinctive element of the pectoral skeleton in birds, which strengthens the shoulder region to withstand the rigor of ...
[4]
Avian cranial kinesis is the result of increased encephalization ...
Mar 17, 2025 · Here, we use three dimensional (3D) muscle modeling and linkage analysis to elucidate the origins of powered cranial kinesis in avian dinosaurs.
[5]
Cretaceous bird with dinosaur skull sheds light on avian cranial ...
Jun 23, 2021 · The known avialan fossils demonstrate that the kinesis in the avian cranium evolved through modifications signified by the non-synchronous loss ...
[6]
Halszkaraptor escuilliei and the evolution of the paravian bauplan
Nov 11, 2019 · ... Paraves, Maniraptora, or Maniraptoriformes in this text. The phylogenetic analysis allowed for the possible plesiomorphic nature of several ...
[7]
Insight into the growth pattern and bone fusion of basal birds ... - PNAS
Oct 9, 2017 · The most distinguishable features are the fusion of the bones in the hand, feet, and pelvis into composite rigid and bony structures.
[8]
Functional constraints on the number and shape of flight feathers
Feb 12, 2024 · Despite the considerable changes in the morphology (three long, clawed manual digits) and proportions of the hand (much longer than the ...
[9]
The origin and early evolution of feathers: implications, uncertainties ...
Feb 19, 2025 · Pennaceous feathers with symmetrical vanes and asymmetrical vanes appeared at the bases of Pennaraptora and Paraves, respectively ...
[10]
Barb geometry of asymmetrical feathers reveals a transitional ...
Each feather vane is composed of a series of branches called barbs attached to the rachis, and each barb consists of a series of smaller branches called ...
[11]
Mosaic evolution in an asymmetrically feathered troodontid dinosaur ...
May 2, 2017 · In having ancestrally asymmetrical vane-widths, the feathers of early paravians (Supplementary Fig. 1) may have been similarly selected upon for ...
[12]
Transformation of the pectoral girdle in pennaraptorans
Feb 29, 2024 · The pectoral girdle, formed by the scapula, coracoid, and furcula, connects the forelimbs and trunk of tetrapods and provides the attachment ...Missing: stabilizing | Show results with:stabilizing<|separator|>
[13]
Wing bone geometry reveals active flight in Archaeopteryx - Nature
Mar 13, 2018 · Its corresponding wingspan, calculated as twice the wing length plus the interglenoid distance, amounts to 0.496 m. From the relation of Witton ...
[14]
Potential for Powered Flight Neared by Most Close Avialan ...
Aug 6, 2020 · Recent studies of early birds and their closest dinosaurian relatives (non-avialan paravian thero- ... ancestral wing loading and specific lift, ...<|separator|>
[15]
Forelimb feathering, soft tissues, skeleton of Microraptor
Jul 1, 2025 · Composite reconstruction of the forewing feathering of Microraptor based on 14 studied specimens. The wing has four to five layers of feathers ...
[16]
Osteohistological analysis of metatarsals reveals new information on ...
Apr 19, 2025 · Additionally, troodontid metatarsals were typically much longer proportionally and express the arctometatarsalian condition, in which metatarsal ...
[17]
Cursorial ecomorphology and temporal patterns in theropod ...
Jan 15, 2025 · Here, we examine temporal evolutionary patterns in the occurrences of cursorial traits such as the arctometatarsus and hind limb proportions.
[18]
Biomechanics of Dromaeosaurid Dinosaur Claws: Application of X ...
Aug 26, 2009 · Dromaeosaurid theropod dinosaurs, such as Velociraptor, possess strongly recurved, hypertrophied and hyperextensible ungual claws on the pes (digit II) and ...
[19]
A new paravian dinosaur from the Late Jurassic of North America ...
Jul 10, 2019 · Paravians are an important radiation of winged coelurosaurs more closely related to birds than to Oviraptor, that include dromaeosaurids, ...
[20]
Keratin Immunoreactivity in the Late Cretaceous Bird Rahonavis ...
... evidence, suggest that original protein- aceous components of keratin claw sheath material may be preserved in the pedal ungual phalanx associated with the.Missing: Paraves | Show results with:Paraves
[21]
Unenlagiinae revisited: dromaeosaurid theropods from South América
The fibula is distally splint-like, similar to that of Buitreraptor (Makovicky et al. 2005). Proximal left tarsals are present (,,), with a ...
[22]
Paravian Phylogeny and the Dinosaur-Bird Transition: An Overview
Synapomorphies possibly exclusive of Anchiornithinae among paravians are dorsal vertebrae with posteriorly expanded neural spines, straight nasal process of the ...
[23]
Testing the function of dromaeosaurid (Dinosauria, Theropoda ... - NIH
Aug 28, 2019 · A musculoskeletal model of the hindlimb and pes of Deinonychus was developed, and mathematical optimization was used to assess the factors that maximize ...
[24]
Cope's rule and the adaptive landscape of dinosaur body size ...
Oct 22, 2017 · Anchiornis huxleyi, Troodontidae, Late Jurassic, 0.58 (0.23–1.4) ... A robust body mass estimate around 70 tonnes was reported using ...
[25]
A Review of Dromaeosaurid Systematics and Paravian Phylogeny
pes of jinfengopterygines, metatarsal III is displaced plantarly at its proximal end relative to the position of metatarsals II and IV (char. 428.1). This ...
[26]
Utahraptor - Utah Friends of Paleontology
Mar 19, 2021 · The largest described Utahraptor ostrommaysorum specimens are estimated to have reached up to 7 metres (23 ft) long and somewhat less than 500 ...
[27]
From dinosaurs to birds: a tail of evolution | EvoDevo - BioMed Central
Jul 29, 2014 · Evolutionary tree of Paraves showing important evolutionary changes. ... avian tails terminating in a fused, distal pygostyle. Considering ...Missing: avialans | Show results with:avialans
[28]
The Evolution of the Theropod Tail into a Stiff Aerodynamic Surface
Between the paravian and avialan nodes, the dorsoventral and lateral joint stiffnesses for the proximal and middle tail decreased (consistent with Hypotheses 2 ...
[29]
Intraskeletal histovariability, allometric growth patterns, and their ...
Jan 10, 2018 · Studying bone histology of extinct paravian dinosaurs (all dinosaurs closer to modern birds than to Oviraptor) is a powerful tool to infer the ...
[30]
How smart was T. rex? Testing claims of exceptional cognition in ...
Apr 26, 2024 · We present revised estimates of encephalization and telencephalic neuron counts in dinosaurs, which we derive from phylogenetically informed ...
[31]
Could Theropod Dinosaurs Have Evolved to a Human Level of ...
Jerison developed a concept called the encephalization quotient (EQ) by which to use brain – body ratios to assess intelligence (Jerison, 1973). The EQ ...
[32]
A new microraptorine theropod from the Jehol Biota and growth in ...
Jan 15, 2020 · The foot is articulated, including both bones of Digit I, which are very reduced and appressed to the lateral surface of Digit II, with the ...
[33]
Intraskeletal Osteohistovariability Reveals Complex Growth ...
Mar 22, 2021 · The endosteal and periosteal layers of slow-growing bone indicate cyclical variation in growth rates. The femur shows regions of coarse ...Missing: intraspecific | Show results with:intraspecific
[34]
[PDF] Osteology oi Deinonychus antirrhopus, an Unusual Theropod from ...
des Deinonychus antirrhopus Ostrom (1969) vor, ein sehr ungewohnlicher, fleisch- fressender Dinosaurier (Ordnung Saurischia, Unterordnung Theropoda) aus der.
[35]
Archaeopteryx and the origin of birds - OSTROM - Wiley Online Library
All available evidence indicates unequivocally that Archaeopteryx evolved from a small coelurosaurian dinosaur and that modern birds are surviving dinosaurian ...Missing: dromaeosaurs | Show results with:dromaeosaurs
[36]
https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1095-8312.1976.tb00244.x
[37]
[PDF] Feathered dinosaurs, flying dinosaurs, crown dinosaurs, and the ...
... (Gauthier and Gall 2001), there is little agree- ment on which clade is to be designated as “Aves.” A precise and universally accepted definition of “Aves ...Missing: Paraves Gallo
[38]
https://repository.si.edu/bitstreams/956cb698-25d0-4a6a-84c3-ca845b63d625/download
[39]
https://doi.org/10.1206/748.1
[40]
https://doi.org/10.1016/j.cub.2015.08.003
[41]
An unusual bird (Theropoda, Avialae) from the Early Cretaceous of ...
Nov 14, 2019 · prima is an avialan: robust pygostyle, robust U-shaped furcula with an interclavicular angle comparable to those of basal birds, strut-like ...
[42]
HIGH RATES OF EVOLUTION PRECEDED THE ORIGIN OF BIRDS
Jan 28, 2014 · The Paraves branch was manually altered from 0 Ma to 500 ... Key patterns are supported here: miniaturization of the Paraves (Turner et al.
[43]
Pre-Archaeopteryx coelurosaurian dinosaurs and their implications ...
Oct 18, 2010 · Fossils, phylogeny, and discovery: A cladistic study of the history of tree topologies and ghost lineage durations. J Vertebr Paleontol ...
[44]
The Solnhofen Limestone of Germany
... Archaeopteryx had feathers is from its exceptional preservation in the Solnhofen Limestone. They also allow more accurate reconstruction of past ecosystems ...Missing: quality | Show results with:quality
[45]
Cranial morphology of Sinornithosaurus millenii Xu et al. 1999 ...
Cranial morphology of Sinornithosaurus millenii Xu et al. 1999 (Dinosauria: Theropoda: Dromaeosauridae) from the Yixian Formation of Liaoning, China · Metrics ...
[46]
Whence the birds: 200 years of dinosaurs, avian antecedents - PMC
Jan 22, 2025 · A rush of discoveries beginning in the early 1990s led to fundamental new insights regarding the evolutionary origins of many aspects of avian ...
[47]
Diuqin lechiguanae gen. et sp. nov., a new unenlagiine (Theropoda
Jun 14, 2024 · Unenlagiines are Gondwanan (Southern Hemisphere) predatory dinosaurs that are nested within Paraves ... There are no synapomorphies in the ...
[48]
A new paravian dinosaur from the Late Jurassic of North ... - PeerJ
Jul 10, 2019 · A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. Research Article.<|control11|><|separator|>
[49]
High-precision 40Ar/39Ar age for the Jehol Biota - ScienceDirect
We obtain an age of 129.7 ± 0.5 Ma for a basaltic lava from the bottom of the Yixian Formation and an age of 122.1 ± 0.3 Ma for a tuff from the lowermost part ...
[50]
Review The Origin and Diversification of Birds - ScienceDirect.com
Oct 5, 2015 · ... pygostyle (a fused, reduced tail bone). These latter birds include ... On the absence of sternal elements in Anchiornis (Paraves) and ...
[51]
Biplane wing planform and flight performance of the feathered ...
A computer simulation of the flight performance of Microraptor suggests that its biplane wings were adapted for undulatory "phugoid" gliding between trees.
[52]
Model tests of gliding with different hindwing configurations in the ...
Jan 25, 2010 · Glide angles ranged from 3° to 21° with a mean estimated equilibrium angle of 13.7°, giving a lift to drag ratio of 4.1:1 and a lift coefficient ...
[53]
Theoretical Aspects of Cursorial Adaptations in Dinosaurs
A theoretical review of the physical constraints on cursorial animals provides a list of the morphological correlates of superior running ability.
[54]
An approach to scoring cursorial limb proportions in carnivorous ...
Jan 27, 2016 · Cursorial-limb-proportion (CLP) scores derived for each of the considered theropod taxa offer a measure of the extent to which a particular species deviates.
[55]
Predictive simulations of running gait reveal a critical dynamic role ...
Sep 22, 2021 · The OCP converged in <5 min, and we obtained a maximum speed of 6.65 m/s, comparable to previous static-based estimates and empirical scaling ...
[56]
Reconstruction of pectoral musculature in non-avialan paravians ...
Oct 23, 2025 · Buitreraptor, Bambiraptor, Archaeopteryx and Confuciusornis do not present a “triosseal canal” because the furcula could contact the acromion ...Missing: loading | Show results with:loading
[57]
[PDF] an arboreal maniraptoran from northeast china
Jul 10, 2005 · While Scansoriopteryx represents an arboreal precursor of Archaeopteryx, in essence it also represents a “proto-maniraptoran”. This suggests ...Missing: prehensile | Show results with:prehensile
[58]
(PDF) Lift devices in the flight of Archaeopteryx - ResearchGate
Aug 6, 2025 · We report on aerodynamic tests using a life-sized model of Archaeopteryx performing in a low turbulence wind tunnel. Our results indicate that ...Missing: finite | Show results with:finite
[59]
A Functional Explanation for Denticulation in Theropod Dinosaur ...
Aug 26, 2009 · The majority of theropod dinosaurs possess ziphodont dentition, which is characterized by laterally compressed, curved, serrated teeth (Langston ...Missing: deinonychosaurs | Show results with:deinonychosaurs
[60]
Developmental and evolutionary novelty in the serrated teeth of ...
Jul 28, 2015 · Ziphodont theropod dinosaurs thus evolved an apomorphic tooth development process that allowed for high serration densities and preserved ...Missing: deinonychosaurs conical
[61]
implications for dietary transition from non-avian dinosaurs to birds
Apr 21, 2020 · We show that avialan teeth are characterized by the presence of simple enamel structures and a lack of porous mantle dentin between the enamel ...
[62]
A Reevaluation of Cooperative Pack Hunting and Gregariousness in ...
Apr 1, 2007 · In this paper we assess the hypothesis of mammal-like cooperative pack hunting in D. antirrhopus and other nonavian theropods by examining the behaviors of ...
[63]
[PDF] article cranial kinesis in dinosaurs: intracranial joints, protractor ...
ABSTRACT—Different forms of intracranial mobility, including streptostyly, pleurokinesis, and prokinesis, have been postulated for many dinosaurs.Missing: troodontids | Show results with:troodontids
[64]
Earliest evidence for fruit consumption and potential seed dispersal ...
Aug 16, 2022 · We show that Jeholornis provides the earliest evidence for fruit consumption in birds, and indicates that birds may have been recruited for seed dispersal.
[65]
Synchrotron imaging of dentition provides insights into the biology of ...
Sep 23, 2016 · ... Hesperornis tooth dentine appears relatively high ... piscivorous marine reptiles, suggesting an association with a piscivorous diet [17].
[66]
Dinosaur biodiversity declined well before the asteroid impact ...
Jun 29, 2021 · We find a negative correlation between the diversity and speciation (sometimes extinction) of a clade, meaning that dinosaur speciation rates ...
[67]
Bird-like characteristics of troodontid theropod eggshell
The presence of two layers (mammillary and squamatic) within the eggshell is either a theropod apomorphy or it arose within Theropoda; the external eggshell ...
[68]
Analysis of dinosaur eggshells: Bird-like Troodon laid 4 to 6 eggs in ...
Apr 3, 2023 · As nests with up to 24 Troodon eggs had been found, the scientists conclude that several Troodon females laid their eggs in communal nests.Missing: papers | Show results with:papers
[69]
Nest and egg clutches of the dinosaur Troodon formosus ... - Nature
Jan 16, 1997 · Nest and egg clutches of the dinosaur Troodon formosus and the evolution of avian reproductive traits. David J. Varricchio,; Frankie Jackson, ...Missing: behavior papers
[70]
A Second Specimen of Citipati Osmolskae Associated With a Nest of ...
Apr 26, 2018 · Here we report on a second nesting oviraptorid specimen (IGM 100/1004) sitting in a brooding position atop a nest of eggs from Ukhaa Tolgod, Omnogov, Mongolia.
[71]
Embryos and eggs for the Cretaceous theropod dinosaur Troodon ...
Aug 10, 2025 · Small (7.5 × 3.5 cm), ellipsoidal eggs with a surface ornamentation consisting of isolated round tubercles represent a new, unnamed ootaxon. The ...
[72]
Fossil bone histology reveals ancient origins for rapid juvenile ...
Nov 28, 2022 · Fibrolamellar bone is generally deposited during the rapid juvenile growth phase of vertebrates with elevated metabolic rates9,50,51,52,53.Missing: Paraves | Show results with:Paraves
[73]
Growth patterns in brooding dinosaurs reveals the timing of sexual ...
Intracortical growth lines in bones from some of the earliest birds suggest that these animals, like their dinosaurian ancestors, took more than 1 year to reach ...<|separator|>
[74]
Sexual dimorphism in dinosaurs - PMC - NIH
Jun 14, 2023 · The results showed that the thigh bones of the dinosaurs displayed subtle differences in shape that were independent of the size of the bone, ...Missing: Velociraptor | Show results with:Velociraptor
[75]
Avian Paternal Care Had Dinosaur Origin - Science
Dec 19, 2008 · Avian Paternal Care Had Dinosaur Origin. David J. Varricchio, Jason R ... View full text|Download PDF · HomeScienceVol. 322, No. 5909Avian ...Missing: text | Show results with:text
[76]
All About Baby Dinosaurs
The young troodontids seem to have been precocial too, as they were born fairly well developed and the nests lack crushed eggshells that would indicate they ...
[77]
An Intermediate Incubation Period and Primitive Brooding in a ...
Aug 20, 2018 · Thus, this tooth age suggests a Troodon incubation period of 74 days, falling midway between avian (44.4 days) and reptilian (107.3 days) values ...