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Pterosaurs: Flying Reptiles of the Mesozoic Darren Naish University of Southampton

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Pterosaurs: Flying Reptiles of the Mesozoic Darren Naish University of Southampton Pterosaurs: Flying Reptiles of the Mesozoic Darren Naish University of Southampton Pterosaurs: winged diapsid rep,les of the Mesozoic, membranous wings supported by super-enlarged 4th finger. Toothed & toothless species. Known since late 1700s; ‘Ptero-dactyle’ named by George Cuvier in 1809. The pterosaur wing Main spar supported by super-enlarged, thickened 4th finger and associated 4th metacarpal with roller-like distal end. Unique, rod-like bone at the wrist - the pteroid – controls posi,on and shape of membrane between shoulder and wrist. Three small, clawed fingers in most (but not all) species. Pteroid is some,mes incredibly long: in Nyctosaurus, more than 50% length of radius Pterosaurs in popular mythology: typically called ‘pterodactyls’, they are scaly, dull-coloured, bat-winged monsters with prehensile feet Pterosaurs: not scaly, but hairy Sordes Excep,onal fossils show that pterosaur bodies were covered in hair-like fibres known as pycnofibres. Best known for Jurassic Sordes from Central Asia, but pycnofibres now known from numerous Triassic, Jurassic and Cretaceous Species from across the family tree. Pterosaurs had fuzzy coats, probably for insulaon. Fibres on top of skull of Sordes Fibres on side of body of anurognathid Jeholopterus The extent of the main part of the wing membrane has been controversial Distal part universally agree to be narrow and slender (= high-aspect), but proximal part argued to be broad, and to incorporate some/all hindlimb, by some, and narrow and aaching to body by others. Fossils show that brachiopatagium did aach to hindlimb, being most extensive in non- pterodactyloids. Brachiopatagium aaching to ,bia in Cretaceous azhdarchoid Brachiopatagium aaching to ankle (perhaps to long 5th toe) in Jurassic rhamphorhynchid Sordes Pterosaur wing membranes were internally complex The ‘darkwing’ Rhamphorhynchus specimen, examined under UV light, reveals the presence of a massive, looping blood vessel system, a layer of subparallel suppor,ng fibres, and a layer with a mesh-like network of connected ,ssue strands. These features suggest that (while thin) the membrane was robust, internally supported, perhaps used in thermoregulaon (or even respiraon). Frey, E., Tischlinger, H., Buchy, M.-C. & Mar,ll, D. M. 2003. New specimens of Pterosauria (Rep,lia) with soe parts with implicaons for pterosaurian anatomy and locomo,on. In Buffetaut, E. & Mazin, J.-M. (eds) Evoluon and Palaeobiology of Pterosaurs. Geological Society Special Publica9on 217. The Geological Society of London, pp. 233-266. Skeletal pneuma>city: pterosaurs, like birds and other saurischian dinosaurs, had pneumac bones. Air-filled sacs connected to the lungs were distributed throughout the body, and occupied most of the skeleton. Demonstrated by pneumac foramina and hollow bones. Pneumac foramina (pf) and hollow bones here illustrated for ornithocheiroid Anhanguera. So->ssue pneumacity: when skeletal pneumacity is present, a series of air-filled sacs connected (by tubes called diver,cula) to the lungs are also present. Pterosaurs had air-sacs throughout the body cavity, also extending along the length of wing and wing-finger. Distribuon of air-sacs in Cretaceous ornithocheiroid Anhanguera Pneumacity present in earliest pterosaurs: was it a primi,ve feature for the group, shared with dinosaurs? Or did it evolve independently? This remains unknown and controversial. What are pterosaurs? They are undoubted members of Neodiapsida, and almost certainly members of Archosauromorpha. But where within this clade do their affini,es lie? S,ll somewhat controversial. Hypothesis 1. Pterosaurs are crown-archosaurs (they have an antorbital fenestra), and share a list of characters with dinosaurs (hinge-like ankle, long neck): there is a pterosaur + dinosaur clade within crown-Archosauria. Hypothesis 2, 3 and 4. Pterosaurs are only convergently similar to dinosaurs, are outside of crown-Archosauria, and (1) are protorosaurs, (2) are close to drepanosaurs, or (3) are close to Archosauria, but not part of it. Specialists are s,ll arguing over these compe,ng hypotheses and pterosaur affini,es have yet to be resolved A quick history of pterosaurs Pterosaurs appear in Late Triassic, persist to end of Late Cretaceous. All early pterosaurs (non-pterodactyloids or “rhamphorhynchoids”) are small predators of small animals. Pterodactyloids evolved in Middle Jurassic: possessed combined nostril + antorbital fenestra (termed nasoantorbital fenestra or NAOF). Evolve giant size (wingspans of c 10 m) and several lineages evolve toothlessness. Pterodactyloids dominate Cretaceous: include oceanic, fish-eang ornithocheiroids, strongly terrestrial dsungaripteroids and azhdarchoids. Only ornithocheiroids and azhdarchoids persist to very end of Late Cretaceous. A quick history of pterosaurs Pterosaurs appear in Late Triassic, persist to end of Late Cretaceous. All early pterosaurs (non-pterodactyloids or “rhamphorhynchoids”) are small predators of small animals. Pterodactyloids evolved in Middle Jurassic: possessed combined nostril + antorbital fenestra (termed nasoantorbital fenestra or NAOF). Evolve giant size (wingspans of c 10 m) and several lineages evolve toothlessness. Pterodactyloids dominate Cretaceous: include oceanic, fish-eang ornithocheiroids, strongly terrestrial dsungaripteroids and azhdarchoids. Only ornithocheiroids and azhdarchoids persist to very end of Late Cretaceous. Non-pterodactyloids (= ‘rhamphorhynchoids’) mostly Triassic and Jurassic, but some lineages persisted into Cretaceous. Generally small (wingspans less than 1 m), generally long-tailed. Short metacarpus, long 5th toes. Curved claws suggest good at climbing. Short hindlimbs. Probably poor terrestrial abili,es. Anurognathids: ,ny size, wide mouths, flexible interphalangeal joints in wing finger Dimorphodonds: long, deep skulls, anterior Unusual in being short-tailed. fang-like teeth. Late Triassic - Early Jurassic. Several Triassic pterosaur taxa (grouped together in Campylognathoididae) are strong heterodont. They have conical anterior teeth suited for grabbing mnd mul,cusped posterior teeth suited for slicing or cung. Oeen extensive tooth wear. Rae9codactylus Upper Triassic, Switzerland. Deep anterior part of mandible, massive nasal horn. Best known form: Eudimorphodon from Italy. Austriadactylus, Late Triassic Austria Eudimorphodon cheek tooth Austriadactylus teeth from (A) premaxilla, (C-D) maxilla & (D-E) posterior part of lower jaw Rhamphorhynchids include taxa with slender jaws and slim, anteriorly poin,ng teeth (like Rhamphorhynchus) as well as taxa with deeper jaws and ver,cally implanted teeth (like Scaphognathus). Rhamphorhynchus-like teeth look good for grabbing fish: this confirmed by stomach contents. But what were scaphognathines doing? Darwinopterus: an evolu,onary intermediate between rhamphorhynchids and pterodactyloids? Long tail, long 5th toe, short metacarpus (like a rhamphorhynchid), but long neck, confluent NAOF like a pterodactyloid. non-pterodactyloid Darwinopterus Modular evolu>on in the origins of pterodactyloids? Darwinopterus suggests that the evolu,onary transi,on from non- pterodactyloids to pterodactyloids occurred in ‘modular’ form (with different parts of the body evolved at different rates). pterodactyloid Skull and neck module seemingly Lü, J., Unwin, D. M., Jin, X., Liu, Y. & Ji, Q. 2009. Evidence for became of pterodactyloid modular evolu,on in a long-tailed pterosaur with a character before the rest of the pterodactyloid skull. Proceedings of the Royal Society B 277, body did. Why? 383-389. Pterodactyloids Evolved during Middle Jurassic from Darwinopterus-like ancestor. NAOF, long & slender pteroid, substan,ally elongate metacarpal IV, short tail, pedal digit V strongly reduced or absent. Ornithocheiroids Toothed and toothless, long-jawed pterodactyloids, some with long, fang-like array of teeth at jaw ,ps probably used for grabbing prey from water. Very small, weakly muscled hindlegs and feet. Strongly specialised for soaring; members of several lineages (pteranodon,ds, nyctosaurids) wholly marine. Pteranodon (and kin): best known ornithocheiroids. Long, gently upcurved, toothless jaws, variable cranial crest. Preserved in Upper Cretaceous marine deposits of American seaway. Marine animals, albatross or frigatebird-like lifestyle. Giant, antler-like crests in Nyctosaurus Suggested by some experts to have formed a windsail-like structure that provided some sort of aerodynamic advantage… Giant, antler-like crests in Nyctosaurus Suggested by some experts to have formed a windsail-like structure that provided some sort of aerodynamic advantage… … but this idea is problemac. Actual shape of bony crest not ‘windsail-like’ at all (it has long posterior spar), bone texture not right for the support of soe- ,ssue membrane, and aerodynamics unlikely to work anyway. Why all the crazy head-crests? Bony crests at the jaw ,ps, above the snout, and on the braincase were common in pterosaurs… ornithocheirid ornithocheiroid tapejarid thalassodromid azhdarchoid azhdarchoid nyctosaurid ornithocheiroid dsungaripterid dsungaripteroid pteranodon,d ornithocheiroid Excep,onally preserved specimens show that soe-,ssue extensions grew on top of some of these bony crests and made them much larger. And some kinds without bony crests (e.g., Pterodactylus) had soe- ,ssue crests that we wouldn’t know about were it not for excep,onal soe-,ssue preservaon. Male Darwinopterus Big cranial crest (also fused pelvis). Female Darwinopterus No cranial crest (also flexible pelvis)… and egg. Darwinopterus: females
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