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Figure 1 in A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs
Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary; dors,dorsal vertebrae; dpc, deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt, metatarsal; po, postorbital; r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3.
Figure S3 in A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs
Figure S3. Histology of the wing phalanx of the new Middle Jurassic pterosaur Dearc sgiathanach, Related to STAR Methods. Thin section in plane polarized light (a), crosspolarized light with a lambda filter (b), cross-polarized light (c), and illustration (d) of histological features: lines of arrested growth (red) and endosteal resorption (blue). Note position of external line of arrested growth very close to the external surface. (e) Inner cortex of wing phalanx under plane polarized light, showing densely vascularized fibrolamellar bone with a line of arrested growth (arrows). (f) Cortex of wing phalanx under cross-polarized light with a lambda filter, showing endosteal lamellae (el), lines of arrested growth (arrows), predominance of primary fibrolamellar bone (flb), and an external zone of parallel-fibered bone (pfb). Note parallel-fibered bone (appearing cyan) adjacent to the line of arrested growth. (g) External cortex of wing phalanx under plane polarized light, showing line of arrested growth (arrow) adjacent to the external bone surface (below dotted yellow line). (h) External cortex of wing phalanx under cross-polarized light, showing parallel-fibered bone (pfb) at the periosteal surface, possibly indicating a decrease in growth rate from the earlier fibrolamellar bone (flb). Arrows denote lines of arrested growth. Abbreviations: el, endosteal lamellae; flb, fibrolamellar bone; pfb, parallel-fibered bone.
Figure 3 in A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs
Figure 3. Postcranial skeleton and dentition of the new Middle Jurassic pterosaur Dearc sgiathanach Photographs of the right manus (A), cervical series (B), pubic region (C), right humerus (D), left humerus (E), left metacarpal-phalanx articular region (F), right maxilla (G), and left pes (H) of NMS G.2021.6.1–4. ac, anterior caudal vertebrae; ar, articular region; c, condyle; cdv, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, dentary; dc, distal condyle; dpc, deltopectoral crest; dvrt, dorsal vertebrae; en, enamel; exp, extensor process; fem, femur; ft, flexor tubercle; gas, gastralia; h, humerus; ic, intercondylar groove; isc, ischium; max, maxilla; mcp, metacarpal; md, maxillary dentition; mg, medial groove; mt, metatarsal; nss, neural spine scar; pa, preacetabular process; poz, postzygapophysis ppb, prepubis; prez, prezygapophysis; rad, radius; rb, rib; sac, sacrum; sm, sesamoid; uc, ulnar crest; ul, ulna; un, ungual; vert, vertebra; wp1, wing phalanx one. Scale, 10 mm per bar. See Figures S2 and S3.
Figure 2 in A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs
Figure 2. Skull of the new Middle Jurassic pterosaur Dearc sgiathanach and comparisons of pterosaur brain and ear endocasts (A–L) Photographs (A–D), segmented MCT scan renderings (E–H), and schematic drawings (I–L) of the skull (NMS G.2021.6.2) in, from top to bottom, left lateral (reversed), dorsal, right lateral, and ventral views. (M–U) Brain and inner ear endocasts of Rhamphorhynchus (based on Witmer et al.7) (M, P, and Q), Dearc (N, R, and S), and Tapejara (based on Eck et al.8) (O, T, and U), shown in the skull in left lateral view and isolated in left lateral and dorsal views, respectively. asc, anterior semi-circular canal; bp, basisphenoid; cbl, cerebellum; cer, cerebrum; ch, choana; chl, channels; cv, third cervical; d, dentary; den, dentition; e, endocast; ec, ectopterygoid; et, edentulous anterior tip; f, frontal (?); hy, hyoid; ipv, interpterygoid vacuity; itf, inferior temporal fenestra; j, jugal; la, lacrimal; md, matrix infilled dent; mx, maxilla; n, nasal; olf, olfactory bulb; opl, optic lobe; p, parietal; pm, premaxilla; po, postorbital; ppd, post-palatal depression; psc, posterior semi-circular canal; ptf, pneumatic foramen; q, quadrate; qj, quadrojugal; rapr, retroarticular process; rh, probable placement of rhamphotheca; sof, suborbital fenestra; sq, squamosal; sbtf, subtemporal fenestra; stf, superior temporal fenestra. In (E)–(L), bones that cannot be distinguished in the MCT scan due to fusion or insufficient resolution are conservatively rendered together in one color. Scale bars, 30 mm. See Figure S2.
Figure S4 in A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs
Figure S4. Results of phylogenetic analysis of the relationships of Dearc sgiathanach and other basal pterosaurs, Related to Figure 4 and STAR Methods. (a) Bootstrap Frequency difference (GC) tree without wildcard or problematic taxa removal from consensus (GC, 100 replicates); (b) Bootstrap absolute frequency tree with problematic taxa removed; (c) Bremer support values, (Bremer.run) with removal of problematic taxa. In each, the red box denotes the clade Angustinaripterini.
Figure S1 in A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs
Figure S1. Locality information for the holotype of the new Middle Jurassic pterosaur Dearc sgiathanach, Related to STAR Methods. Map and geological context showing: Isle of Skye (a); geological map of the locality where the specimen was found, Brothers' Point (b); stratigraphic section at the discovery site (c); orthophoto of the pterosaur fossil location in relation to dinosaur trackways and evidence of desiccation (d). Figures c) & d) modified after dePolo et al. (2020).
Figure S2 in A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs
Figure S2. Skull and anterior cervical vertebrae of the new Middle Jurassic pterosaur Dearc sgiathanach visualised by X-ray computed microtomography (manual & machine learning segmentation), Related to Figures 1-3. Manually segmented skull in: (a) dorsal; (b) ventral; (c) right lateral; (d) left lateral; (e) anterior; (f) posterior views. Colour code: yellow – palate & pterygoid; purple – dentary; black – teeth; green – fused dorsal cranial elements; blue – basipterygoid; light blue – dorsal unfused cranial bones. Manually segmented atlas, axis & anterior cervical complex in: (g) lateral; (h) ventral; (i) dorsal views. Cervical close up with schematic line art in: (j) dorsal and (k) ventral views. Colour code: dark yellow – 3rd cervical; cream – atlas & axis. Manually segmented brain and inner ear endocast in: (l) anterior; (m) dorsal; (n) ventral; (o) left lateral views. Colour code [annotated in the figure] channels; dark blue – optic lobes; pink – flocculus; purple – cerebrum & cerebellum; light blue – cranial nerves; WEKA trained machine learning segmentation results (p) in dorsal and ventral (left to right, respectively). Line art of select elements (q) cervical vertebral series in dorsal view, abbreviations: cv# - cervical vertebra; cr - cervical ribs; ns - neural spine; NMS G.2021.6.3 counterslab (r) in ventral orientation, abbreviations: lcv - long caudal vertebra; scv - short caudal vertebra; isc - ischium; ppb - prepubis; sv - sacral vertebra (?); gas - gastralia; sac - sacrum; dv - dorsal vertebra; rib - dorsal ribs; hum - humerus; sc - scapula coracoid; and dorsal view of right humerus (s); abbreviations: hum - humerus; ra - radius; ul - ulna. Comparative size diagram of select non-pterodactyloid humeri (t) [annotated in image]. Scales bars=20 m
Figure 4 in A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs
Figure 4. Phylogenetic relationships of the Middle Jurassic pterosaur Dearc sgiathanach and wingspan estimates for Jurassic pterosaurs Strict consensus of most parsimonious trees from phylogenetic analysis, with silhouettes scaled to wingspan (Dearc = ca. 2.0 m) (1), and skull reconstructions of key taxa: (A) Scaphognathus crassirostris (based on GPIB 1304), (B) Dorygnathus banthensis (based on SMNS 55886), (C) Rhamphorhynchus muensteri (based on NHMUK R 37002), (D) Angustinaripterus longicephalus (based on ZDM T8001), and (E) Dearc sgiathanach. Wingspan plot (2): estimated wingspan range for isolated pterosaur remains from the Taynton Limestone collection. See Data S1D for complete element list. (A) NHMUK PV R 36634, (B) GSM 113726, (C) OUM J28352, (D) NHMUK PV R 38016, (E) NHMUK PV R1362, (F) LL12158, (G) NHMUK PV R 40126, (H) NMS G.2021.6.1-4 (Dearc), (I) OUM J23046, (J) OUM J28273, (K) NHM UK PV R 40126 B, (L) OUM J28319, (M) OUM J28307, (N) OUM J28271, (O) OUM J28354, (P) LL12160, (Q) OUM J23047, and (R) MJM L K1995. See Figure S4 and Data S1 and S2.
Fig. 10 in An exceptionally preserved Sphenodon-like sphenodontian reveals deep time conservation of the tuatara skeleton and ontogeny
Fig. 10 Evolutionary changes on the temporal region of sphenodontians. Many early-evolving lepidosaurs retain a single temporal fenestration as the jugal (cyan) does not contact the quadrate/quadratojugal (green) posteriorly. Afull development of the lower temporal bar and double temporal fenestration evolved independently at least twice in sphenodontians, once among clevosaurids and once in sphenodontines—an adaptation for stabilizing the quadrate and reducing overall stress in the skull during hard biting58. The latter is unique among lepidosaurs by including contributions from both the jugal and quadratojugal—a morphology convergent with many non-lepidosaurian early diapsid reptiles. Red circle, complete lower temporal bar; blue circle: incomplete lower temporal bar; blue-red gradient circle, lower temporal bar incomplete in juveniles but complete in adults. Skull drawings from top to bottom: Gephyrosaurus (drawn by TRS based on ref. 20), Megachirella (re-drawn by TRS from ref. 33), Diphydontosaurus (drawn by TRS based on ref. 20), Clevosaurus (drawn by TRS based on ref. 20), Palaeopleurosaurus (drawn by TRS based on ref. 20), Navajosphenodon (drawn by A. Brum), and Sphenodon (drawn by TRS based on MCZ R4702).
Fig. 9 in An exceptionally preserved Sphenodon-like sphenodontian reveals deep time conservation of the tuatara skeleton and ontogeny
Fig. 9 Phylomorphospace of early lepidosaurs and sphenodontians using discrete morphological characters. Clade "A" refers to the clade recovered by the final analysis (relaxed morphological clock Bayesian inference) both here and in Simões, et al.19, including Homeosaurus, pleurosaurids, and saphaeosaurids. For figures with individual taxon names, see Supplementary Fig. 4. Cyno Cynosphenodon, Kaw Kawasphenodon, Nav Navajosphenodon, Sphe. Sphenodon, Sphf. Sphenofontis.
Fig. 8 in An exceptionally preserved Sphenodon-like sphenodontian reveals deep time conservation of the tuatara skeleton and ontogeny
Fig. 8 Majority rule consensus tree from the relaxed morphological clock Bayesian inference analysis with tip dating. Results indicate the phylogenetic relationships among sphenodontians highlighting the placement of N. sani (in bold), clade posterior probabilities (top node values in bold), and median divergence times (bottom node values). Purple node error bars represent the 95% highest posterior density estimates for divergence times. Skull illustrations (all photos taken by TRS) for each major clade are, from top to bottom, Megachirella wachtleri (Squamata), Clevosaurus brasiliensis (Clevosauridae), Homeosaurus maximiliani (Homeosaurinae), Pleurosaurus gingsburi (Pleurosauridae), Kallimodon pulchellus, Priosphenodon avelasi (Eilenodontinae), and Sphenodon punctatus (Sphenodontinae).
Fig. 7 in An exceptionally preserved Sphenodon-like sphenodontian reveals deep time conservation of the tuatara skeleton and ontogeny
Fig. 7 Postcranial skeleton of the holotype of N. sani (MNA.V.12442). a Atlas, axis, and cervical vertebrae 3 and 4 in ventral view. b Part of Atlas–axis complex in dorsal view. c Dorsal vertebrae in ventral view. d Caudal vertebrae in lateral view. e Pelvic girdle and pygals in ventrolateral view. f Right pes in partial articulation. g Left pes with only distal phalanges preserved. h Left scapula and coracoid in lateral view. i Right(?) pubis and ilium. j Left forearm in posterior view. k Left forearm in anterior view. l Right forearm in anterior view (slightly displaced radius and ulna). m Right forearm in posterior view. Ace acetabulum, As astragulus, Ax axis, Ax.Ce. axis pleurocentrum, Ax.NS. axis neural spine, At.NA. atlas neural arches, Ca.V. caudal vertebra, Co coracoid, Ce.V. cervical vertebra, Dp.Cr. deltopectoral crest, Ent.Fr. entepicondylar foramen, H humerus, H.H. humeral head, Il ilium, Il.Bl. iliac blade (partially preserved), Is isquium, N.S. neural spine, Obt.Fr. obturator foramen, Od. odontoid (atlas centrum), Ol.Pr. olecranon process, Pn.Ph. penultimate phalanx, Po.No. posterior notch on acetabulum, Pu pubis, Pu.Pr. anterior pubic process, Py pygals, Ra radius, Ra.Cd. radial condyle, Sca scapula, Ul ulna, V.Cr. midventral crest, I–V digit number. Scale bars = 1 mm.
Fig. 6 in An exceptionally preserved Sphenodon-like sphenodontian reveals deep time conservation of the tuatara skeleton and ontogeny
Fig. 6 Conservation of ontogenetic stages from Navajosphenodon to Sphenodon. Ontogenetic series on the maxilla (a–c) and dentary (d–f) of Sphenodon punctatus (rescaled to the same length). Ontogenetic series on the maxilla (g–i) and dentary (j–l) of Navajosphenodon sani. Specimen numbers: FMNH 207433 (b, e); FMNH 11113 (c, f); MCZ VP VP 9094 (g); MCZ VP VP 9100 (h); MCZ VP VP 9093 (i); MCZ VP VP 101564 (top left), MCZ VP VP 9094 (top right), MCZ VP VP 9099 (bottom left), MCZ VP VP 101569 (bottom right) (j); MNA.V.12442 (k); MCZ VP 9093 (l). a, d Re-drawn from ref. 43. Add.T. additional teeth, Alt.T. alternating teeth, Ang angular, D.C.Pr. dentary coronoid process, D.Po.Pr. dentary posterior process, Ed. edentulous region, Hat.T. Hatchling teeth, M.Pr. mentonian process, Me.C. Meckelian canal, Suc.T. Successional teeth, Worn worn out teeth. Scale bars = 1 mm.
Fig. 5 in An exceptionally preserved Sphenodon-like sphenodontian reveals deep time conservation of the tuatara skeleton and ontogeny
Fig. 5 Mandibles of the holotype of N. sani (MNA.V.12442). Left mandible in lateral view (a) and medial view (b). Right dentary in medial view (c). Left post-dentary bones in dorsal view (d). Ce.Rd. central ridge, Gl. glenoid articulation, Me.C. Mekelian canal, RAP retroarticular process, San.C.Pr. surangular coronoid process, Sym. symphysis, V.Cr. ventral crest. Scale bars = 10 mm (a, b) and 1 mm (c, d).
Fig. 4 in An exceptionally preserved Sphenodon-like sphenodontian reveals deep time conservation of the tuatara skeleton and ontogeny
Fig. 4 Palatal region and braincase of the holotype of N. sani (MNA.V.12442). Palatal region in dorsal (a) and ventral (b) views. Basisphenoid in ventral (c), dorsal (d), anterior (e), and right lateral (f) views. Arc.Fl. Arcuate flanges, Bpt.Pr. basipterygoid process, Ca.Op. openings for internal carotid arteries, Cl. Pr. clinoid process, Cu.Pr. cultriform process, Do.Se. dorsal sella, L.Pr. lateral process, Pal.Asc.Pr. palatine ascending process, Pal.T. palatine teeth, Ptg.T. pterygoid teeth, Q.Pr. quadrate process, Sel.Tu. sella turcica, Tr.Cr. trabeculae cranii, VI passage for cranial nerve VI (abducens canal). Scale bars = 1 mm.
Fig. 2 in An exceptionally preserved Sphenodon-like sphenodontian reveals deep time conservation of the tuatara skeleton and ontogeny
Fig. 2 Micro CT-scanned and fully segmented skull and mandibles of the holotype of N. sani (MNA.V.12442). a Right ventrolateral view. b Left dorsolateral view. Art articular, Boc basioccipital, Bsp basisphenoid, C coronoid, CbI first ceratobranchial, D dentary, Ect ectopterygoid, Epi epipterygoid, F frontal, J jugal, M maxilla, N nasal, P parietal, Pal palatine, PFr postfrontal, PM premaxilla, Po postorbital, Pra prearticular, PrF prefrontal, Ptg pterygoid, Q-Qj quadrate-quadratojugal, San surangular, Spm septomaxilla, Sq squamosal, (l) left side and (r) right side. Scale bar = 1 mm.
Fig. 3 in An exceptionally preserved Sphenodon-like sphenodontian reveals deep time conservation of the tuatara skeleton and ontogeny
Fig. 3 Individual elements of the skull of the holotype of N. sani (MNA.V.12442). a Premaxilla in lateral (left) and posterior (right) views. b Left maxilla in lateral (left) and medial (right) views. c Left maxilla in occlusal view. d Nasals in dorsal view. e Left prefrontal in lateral (left) and medial (right) views. f Right septomaxilla in ventral view. g Right postfrontal in lateral (left) and medial (right) views. h Right jugal and associated quadratojugal in lateral view. i Fused left quadrate–qudratojugal complex in posterior view. j Left jugal in lateral view. k Right postorbital in medial view. l Left postorbital and squamosal in lateral view. m Frontals and parietals in dorsal view. n Frontals in ventral view. o Right frontal in lateral view. Relative bone positions in h, l, m are as preserved in the specimen. Ant.Pr. anterior process, Av.Pr. anteroventral process, Acr.T. acrodont teeth, Ant.Pr. anterior process, AnL.Pr. anterolateral process, br. broken edge, D.Pr. dorsal process, Di.Pr. distal process, E.N. external nares, J.Ft. facet for jugal, N.Pr. nasal process, P.Pr. posterior process, Pal.Ft. facet for maxillary process of palatine, PFr.Ft. facet for postfrontal, Pm.Cr. posteromedial crest, Pm.Pr. premaxillary process, Po.Ft. facet for postorbital, Po.Pr. postorbital process, PoL.Pr. posterolateral process, PrF.Ft. facet for prefrontal, Pv.Pr. posteroventral process, Q quadrate, Qj quadratojugal, Qj.Fr. quadratojugal foramen, Sof.Pr. subolfactory process, St.Pr. supratemporal process, T.Ap. tooth apex, V.Pr. ventral process, Vl.Pr. ventrolateral process, Vo.Ft. facet for vomeronasal organ. Scale bars = 1 mm.
FIG. 27 in Skeleton of Early Miocene Bathyergoides neotertiarius Stromer, 1923 (Rodentia, Mammalia) from Namibia: behavioural implication
FIG. 27. — Femora of extant fossorial rodents and GT 50'06: Cryptomys hottentotus (Lesson, 1826) AZ 834, right femur; A, anterior view; F, posterior view, K, medial view; Bathyergus janetta Thomas & Schwann, 1904 TM 39332, left femur; B, anterior view; G, posterior view; L, medial view; Heliophobius argentocinereus Peters, 1846 TM 45931, left femur; C, anterior view; H, posterior view; M, medial view; Tachyoryctes splendens (Rüppell, 1835) 820 38 M 1, right femur; D, anterior view; I, posterior view; N, medial view; Bathyergoides neotertiarius Stromer, 1923 GT 50'06, left femur E, anterior view; J, posterior view; O, medial view. Scale bars: 1 cm.
FIG. 18 in Skeleton of Early Miocene Bathyergoides neotertiarius Stromer, 1923 (Rodentia, Mammalia) from Namibia: behavioural implication
FIG. 18. — Left calcaneum of GT 50'06: A, dorsal view; B, lateral view; C, plantar view; D, medial view; E, distal view; F, Proximal view. Scale bar: 1 cm.
FIG. 24 in Skeleton of Early Miocene Bathyergoides neotertiarius Stromer, 1923 (Rodentia, Mammalia) from Namibia: behavioural implication
FIG. 24. — Mandible of extant fossorial rodents and GT 50'06; A-C, ventral view; A, Heliophobius argentocinereus Peters, 1846 TM 45931; B, Cryptomys hottentotus (Lesson, 1826) AZ 834; C; Bathyergus janetta Thomas & Schwann, 1904 TM 39332; D-F, left lateral view; D, Heliophobius argentocinereus Peters, 1846 TM 45931; E, Cryptomys hottentotus (Lesson, 1826) AZ 834; F, Bathyergus janetta Thomas & Schwann, 1904 TM 39332; G-I, Bathyergoides neotertiarius Stromer, 1923 GT 50'06; G, left lateral view; H, right lateral view; I, ventral view. Scale bars: 1 cm.
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