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Extended Data Fig. 5 in A new Jurassic scansoriopterygid and the loss of membranous wings in theropod dinosaurs

Extended Data Fig. 5 | Comparisons of hand morphology among Scansoriopterygidae. a–c, Line drawings of the hands of Ambopteryx (a), Epidendrosaurus (b) and Yi (c). Scale bars, 10 mm (a, b), 20 mm (c).

Extended Data Fig. 4 in A new Jurassic scansoriopterygid and the loss of membranous wings in theropod dinosaurs

Extended Data Fig. 4 | Forelimb comparisons between Ambopteryx and Yi. a, Left forelimb of Ambopteryx (IVPP V24192). b–d, Left forelimb (b), right humerus (c) and the right styliform element (d) of Yi (STM 31-2). The proximal margins of the humeri are marked in white dashed lines to show the differences between these two taxa. Abbreviations as in Figs. 1, 2, except for: st, styliform element. Scale bars, 10 mm (a), 20 mm (b–d).

Extended Data Fig. 3 in A new Jurassic scansoriopterygid and the loss of membranous wings in theropod dinosaurs

Extended Data Fig. 3 | Anatomy of Ambopteryx, IVPP V24192. a–d, Interpretative drawings of the neck and pectoral girdle (a), caudal vertebrae and pygostyle (b), left forelimb (c) and pelvis and left hindlimb (d). Abbreviations as in Figs. 1, 2, except for: ca, caudal vertebrae; cm, calcaneum; de, deltopectoral crest of humerus; do, dorsal vertebrae; I–III, metacarpals I–III; ip, iliac peduncle of ilium; mtII–IV, metatarsals II–IV; p1–4, pedal digits I to IV; and st, styliform element. Scale bars, 10 mm (a–d).

Fig. 1 in A new Jurassic scansoriopterygid and the loss of membranous wings in theropod dinosaurs

Fig. 1 | Holotype of A. longibrachium, IVPP V24192. a, b, Photograph (a) and line drawing (b) of the holotype of A. longibrachium (IVPP V24192). as, astragalus; ca, calcaneum; ce, cervical vertebrae; cv, caudal vertebrae; dr, dorsal rib; dv, dorsal vertebrae; ga, gastroliths; gs, gastralia; is, ischium; lc, left coracoid; lf, left femur; lfi, left fibula; lh, left humerus; li, left ilium; lm2, lm3 and lm4, left metatarsals II, III and IV; lmd1, lmd2 and lmd3, left manual digits I, II and III; lr, left radius; ls, left scapula; lt, left tibia; lu, left ulna; mc1, mc2 and mc3, metacarpals I, II and III; nf, neck feathers; pd1, pd2, pd3 and pd4, left pedal digits I, II, III and IV; pu, pubis; py, pygostyle; rc, right coracoid; rh, right humerus; ri, right ilium; rmd3, right manual digit III; rr, right radius; rra, right radiale; rs, right scapula; ru, right ulna; rua, right ulnare; se, styliform element; sk, skull; sv, sacral vertebrae; to, tooth. Brown and dark grey shadings denote feathers and membranous tissues, respectively; blue and light blue shadings indicate the potential bony stomach content; light grey denotes unidentified matrix. Red box indicates the location from which the sample was taken for histological analysis. Scale bars, 10 mm.

Extended Data Figure 3 in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Extended Data Figure 3 | Close-ups of the three counter slab fragments and corresponding slab regions of STM 31-2. Top row: isolated piece from counter slab preserving cranial region (left), and corresponding region of slab (right); middle row: isolated piece from counter slab preserving forelimb region (left), and corresponding region of slab (right); bottom row: isolated piece from counter slab preserving hindlimb region (left), and corresponding region of slab (right).

Extended Data Figure 1 in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Extended Data Figure 1 | Fossil-bearing beds of the Middle–Upper Jurassic Tiaojishan Formation in Qinglong County, Hebei. The Mutoudeng Locality (top) and Gangou Locality (bottom) are geographically close to each other (approximately 20 km apart) and very similar in their sedimentology and fossil content. The bottom image shows an excavation we organized at the Gangou Locality.

Extended Data Figure 8 in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Extended Data Figure 8 | Scanning electron microscopy sample locations. Numbered circles indicate the locations of samples from the slab and counter slab. 1, distal part of cranial feathers; 2, middle part of tibial feathers; 3, distal part of humeral feathers; 4, distal part of ulnar feathers; 5, proximal part of cranial feathers; 6, proximal part of neck feathers; 7, distal part of humeral feathers; 8, distal part of humeral feathers; 9 and 10, membranous soft tissue near digit II; 11, membranous soft tissue near digit IV; 12, middle part of tibial feathers.

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Theropod Toe Bone

A toe bone of a theropod from Morocco. Possibly from a spinosaurd phalanx III from toe 4. Location: Kem Kem Beds, Morocco Age: Cenomanian, Late Cretaceous Length: 2.9 cm Source: Objaverse 1.0 / Sketchfab

Fig. 4 in The Jordan theropod (Maastrichtian, Montana, U.S.A.) referred to the genus Aublysodon.

Fig. 4 - Comparison of The skulls of A, Aubtysodon; B, Coelopbysis (AMNB 7224); C, Dromaeosaurus (modified from Coibert & Russell, 1969, Fig. 1); D, Velociraptor (modified from Barsbold, 1982, Fig. la); E, Sauromitboides (modified from Barsbold, 1974, Pl. 1), V, Dromiceiomimus (modified from Russell, 1972, Fig. 5, and ROM 840); G, Caegnatbus (modified from Sternberg, 1949, Fig. 1, and Barsbold, 1982, Fig. c); H, Albeltosaurus libratus (modified from Gilmore, 1946, Pl. 4, Fig. 3); I, Daspletosaurus torosus (modified from Russell, 1970, Fig. 6); J, " Albertosaurus " lancensis (modified from Gilmore, 1946, Pl. 1); H, Tyrannosaurus (modified from Osborn, 1912, Fig. 1); L, lingual (top) and labial (bottom) views of the dentaries of " Cbirostenotes " (modified from Gilmore, 1924, Fig. 3). Scale bar represents 10 cm. Comparaison des cranes des divers théropodes: A, Aubtysodon; B, Coelopbysis (AMNB 7224); C, Dromaeosaurus (modifiés d'après Colbert & Russell 1969, Fig. 1); D, Velociraptor (modifié d'après Barsbold 1982, Fig. la); E, Sauromitboides (modifié d'après Barsbold 1974, Pl. 1); F, Dromiceiomimus (modifié d'après Russell 1972, Fig. 5 et ROM 840); C, Caenagnatbus (modifié d'après Sternberg 1949, Fig. 1 et Barsbold 1982, Fig. c); H, Albertosaurus libratus (modifié d'après Gilmore 1946, PI. 4, Fig. 3); I, Daspletosaurus torosus (modifié d'après Russell 1970, Fig. 6); J, " Albertosaurud' lancensis (modifié d'après Gilmore 1946, Pl. 1); K, Tyrannosaurus (modifié d'après Osborn 1912, Fig. 1); L, les vues linguale (en haut) et labiale (en bas) des dentaires du " Cbirostenotes " (modifié d'après Gilmore 1924, Fig. 3). Echelle = 10 cm.

Fig. 2 in The Jordan theropod (Maastrichtian, Montana, U.S.A.) referred to the genus Aublysodon.

Fig. 2 - Cheek teeth of Aubtysodon mirandus. Left sixth maxillary tooth, as preserved, in lateral (A) and posterior (B) aspects. Right fourth dentary tooth, rendered in unworn condition, in anterior (C) and labial (D) aspects. Scale bar represents 10 mm. Les dents jugales de VAubtysodon mirandus. Sixième dent maxiliaire gauche telle qu'elle est conservée en vue laterale (A) et vue postérieure (B). Quatrième dent du dentaire droit, figurées sans usure en vue antérieure (C) et en vue labiale (D). Echelle: 10 mm.

Fig. 2 in The Jordan theropod (Maastrichtian, Montana, U.S.A.) referred to the genus Aublysodon.

Fig. 4 in The Jordan theropod (Maastrichtian, Montana, U.S.A.) referred to the genus Aublysodon.

Extended Data Figure 9 in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Extended Data Figure 9 | Scanning electron microscopy images of melanosomes and melanosome impressions preserved in Yi qi holotype (STM 31-2). a–m, Note the high morphological diversity of the preserved melanosomes in the feathers, which include: densely distributed, small round phaeomelanosomes and sparsely distributed, medium-sized oval eumelanosomes in the feathers near the skull (a); medium-sized elongate elliptical eumelanosomes in the feathers near the skull (b, c), neck (d, e), humerus (f, g), and ulna (h, i, j); and large oval and elliptical eumelanosomes in the feathers near the tibiotarsus (k, l). The subspherical phaeomelanosomes in the sheet-like soft tissue (m) appear to be less densely distributed than the melanosomes in the feathers.

Extended Data Figure 7 in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Extended Data Figure 7 | Aerodynamic apparatuses in Yi and other tetrapods. a–c, Wing of Yi, Bat Model (a), Maniraptoran Model (b), and Frog Model (c); d, bat wing; e, pigeon wing; f, pterosaur wing; g, giant Japanese flying squirrel wing. Yellow colour indicates styliform element in a, b, c, d, f, and g.

Figure 3 in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Figure 3 | Simplified coelurosaurian phylogeny showing the recovered position of Yi. The skeletal silhouette and two possible alternative planform reconstructions of Yi highlight the proportionally long and robust forelimbs and large leg feathers that Yi shares with other basal paravian theropods, indicating the presence of aerial capability, and the inferred membranous wings, a feature unique among known paravians but seen in most other gliding or flying tetrapods. Various uncertainties, such as how the styliform element is oriented and whether membranous tissue is present lateral to the trunk as in most volant tetrapods, imply that a variety of reconstructions of the aerodynamic apparatus of Yi are currently plausible (see Supplementary Information for additional possible reconstructions).

Extended Data Figure 6 in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Extended Data Figure 6 | Elemental compositions of the styliform element and comparison samples based on EDS analyses. EDS spectra derived from: a, right manual phalanx II-1; b, right styliform element; c, feathers associated with right tibia; and d, sedimentary matrix.

Figure 2 in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Figure 2 | Soft tissues preserved in Yi qi holotype (STM 31-2). a–e, Feathers above the skull (a), along the humerus (b) and along the tibiotarsus (c); isolated basally converging (d) and brush-like (e) feathers; f, sheet-like soft tissue associated with the right forelimb (yellow arrows point to patches of sheet-like soft tissue); g, comparison of sheet-like soft tissue (above digit II) and individual feathers (below digit II).

Figure 1 in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Figure 1 | Yi qi holotype (STM 31-2). a, b, Photograph (a) and line drawing (b) of specimen; c, skull and mandible in lateral view; d, premaxillary tooth in lateral view; e, left manus; f, styliform elements (the distally unexposed left styliform element articulates with the wrist, and the orientation of the right styliform element implies a similar relationship to the carpus even though its proximal part is missing). Light and dark grey shading indicates feathers and membranous tissues, respectively. an, angular; cv, cervical vertebrae; d, dentary; dr, dorsal ribs; emf, external mandibular fenestra; en, external naris; f, frontal; lf, left femur; lh, left humerus; lmd2–4, left manual digits 2–4; lmt, left metatarsals; lr, left radius; ls, left scapula; lse, left styliform element; lu, left ulna; mb, mandible; mcII–IV, metacarpals II–IV; n, nasal; or, orbit; p, parietal; phII1 to IV4, phalanges II-1 to IV-4; pma, premaxilla; rmd2–4, right manual digits 2–4; rf, right femur; rfi, right fibula; rh, right humerus; rmc, right metacarpals; rmt, right metatarsals; rr, right radius; rse, right styliform element; rt, right tibiotarsus; ru, right ulna; sk, skull. Scale bar, 2 cm.

cranial feathers; 6, proximal part of neck feathers; 7, distal part of humeral feathers; 8, distal part of humeral feathers; 9 and 10, membranous soft tissue near digit II; 11, membranous soft tissue near digit IV; 12, middle part of tibial feathers. in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

cranial feathers; 6, proximal part of neck feathers; 7, distal part of humeral feathers; 8, distal part of humeral feathers; 9 and 10, membranous soft tissue near digit II; 11, membranous soft tissue near digit IV; 12, middle part of tibial feathers.

elliptical eumelanosomes in the feathers near the skull (b, c), neck (d, e), humerus (f, g), and ulna (h, i, j); and large oval and elliptical eumelanosomes in the feathers near the tibiotarsus (k, l). The subspherical phaeomelanosomes in the sheet-like soft tissue (m) appear to be less densely distributed than the melanosomes in the feathers. in A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

elliptical eumelanosomes in the feathers near the skull (b, c), neck (d, e), humerus (f, g), and ulna (h, i, j); and large oval and elliptical eumelanosomes in the feathers near the tibiotarsus (k, l). The subspherical phaeomelanosomes in the sheet-like soft tissue (m) appear to be less densely distributed than the melanosomes in the feathers.