Index

Page numbers in italics refer to Figures. Page numbers in bold refer to Tables. acetabulum, orientation 45 bite marks on pterosaurs 15–16 Adriatic Carbonate Platform 210 biting, induced bending moments 25 Aerodactylus spp. 125, 127 Blackhawk Formation 11,12 A. scolopaciceps 130 body mass, effect on predation 19 aerodynamics 57 bone aktinifibrils 57 evolution for flight 69 Altmuehlopterus rhamphastinus 140 textures 70–71 Angustinaripterus spp. 31, 37, 184 see also jaw bone A. longicephalus 27, 32, 34, 35, 36, 40, 41 Brazil Anhanguera spp. 35, 52 Crato Formation 13, 195, 196 A. piscator 46, 228 Lagersta¨tten 1 A. santanae 26, 27, 29, 33, 36, 39, 40, 41,46 Santana Formation 14,16 antorbital fenestae 25 A. santanae 26, 27, 29, 33, 36, 36, 39, 40, 41,46 Anurognathidae 95 Breakfast Bench Facies (Wyoming) 105, 106 Anurognathus spp. 30, 31, 32, 42, 95, 103 Buckland, William 1 A. ammoni 27, 32, 34, 35, 36, 40, 41 Arambourgiania philadelphiae 159, 167 Cambridge Greensand Formation 17 descriptions of possible samples 161–167 Campylognathoides spp. 30, 31, 42, 176 evaluation 167–168 C. liasicus 27, 32, 34, 35, 36, 40, 41 systematics 160–161 Canada Araripe Basin (Brazil) 195, 196 Dinosaur Park Formation 14,16 Araripesaurus santanae 228 carpal 100 Ardeadactylus longicollum 130, 130 Caulkicephalus trimicrodon 221, 223 Argentina Caviramus spp. 176 Lagarcito Formation 10 Cerro del Pueblo Formation (Mexico) 11,12 embryo discoveries 83, 84, 85, 86, 87, 88 cervical column 201 comparison with other sites 92 Chaoyangopteridoidae 201, 203, 205 maturity criteria 91–92 cheiropatagium 58 measurements 87 China preservation 84–86 embryo studies 83 skeletal elements comparison with Argentina embryo 92 limbs 89–90 Jehol Group 195 pectoral girdle 89 Junggar Basin 149 skull 86–87 Jiufotang Formation 95 vertebral column 87–89 Vesperopterylus lamadongensis 95–96 taxonomic assignment 90–91 comparative anatomy 101, 103 Atherfield Clay Formation 46 description 96, 100–101 Aurorazhdarchidae 135 measurements 100 Aurorazhdarcho micronyx 130 palaeoenvironment 101 Austriadactylus 120 phylogeny 102, 103 Australia, Toolebuc Formation 14,16 skeleton 96, 98, 99 azhdarchids 12, 20, 46, 47, 112, 113, 159, 160–161, 181, Lagersta¨tten 1 201, 204, 216 clade retention index (CRI) 132, 133–136, 136 Azhdarcho spp. 115 cladistics 131–133, 133–136 A. lancicollis 160, 216, 228 Colobrorhynchus spp. 221, 223 C. clavirostris 221, 223 Balqa Group 160, 161 Comodactylus ostromi 105 Batrachognathus spp. 95, 103 comparative anatomy, Diopecephalus 136–137 Bavarian State Collection for Palaeontology and condyles 215 Geology (Munich) 159, 168 consistency fork index (CFI) 132 bending moments, impact on skull 25 coprolites 10–11 Bexhill coracoid 97 geological setting 222, 222 Crato Formation (Brazil) 13, 195, 196 new pterosaur find 222, 224, 224 Cretaceous specimens 159, 187, 209 assignment and classification 229, 231 Australia 14,16 skeletal characteristics 225–229 Brazil 195 systematics 224–225 embryos 83, 84 bipedal gait 45 extreme bone lightness 69 234 INDEX

Cretaceous specimens (Continued) limbs 89–90 Jordan 159 pectoral girdle 89 Er Ruseifa old phosphate mines 160, 161 skull 86–87 Thailand 181 vertebral column 87–89 trackways 11,12 taxonomic assignment 90–91 Wealden Supergroup 221, 223 Eudimorphodon spp. 28, 30, 30, 31, 38, 42, 120, 176 Wight, Isle of 46 E. ranzii 8, 9, 27, 32, 34, 35, 36, 39, 40, 41 Vectidraco daisymorrisae muscle study 46 Eurazhdarcho langendorfensis 16 methods 47 results 47–51 fangs 108, 110, 111, 116–118, 117, 120 results discussed 51–54 feeding traces 11 crucopatagium 58 femur studies Ctenochasma spp. 31, 31–32, 34, 36, 42 Arambourgiania 163, 165–166 C. elegans 130, 130 Dsungaripteroidea 107, 108, 111–114 113, 114 C. gracile 27, 33, 36, 40, 41 embryo 87 Cuba 184 Rhamphorhynchus 173, 174, 177, 177 Cuvier, Georges 1, 45 Vesperopterylus 101 Cycnorhamphus spp. 42, 130 Ferron Sandstone (Utah) 187 C. suevicus 26, 27, 33, 36, 40, 41, 130, 130 fibula 101 fish, as food 8, 10 Daitingopterus rhamphastinus 141 flight, bone evolution for 69 Darwinopterus spp. 31, 83 Flugsaurier symposia 3 Del Monte Mines (Utah) 11 flutter Dendrorhynchoides spp. 95, 96 problems of 57 Dermodactylus montanus 105 limits to membrane tension 63–64 digital modelling of skull food webs, role of pterosaurs 7, 19–20 methods foraging behaviour geometry 26–27 difficulties in proving 12–13, 15 orbital area 30 evidence for 7 stiffness 27–30 gut contents 8–10 results 30–31 regurgitates 10 orbit 35–36, 40 trace fossils 12 stiffness 31 Franconia laminated limestone 125, 130, 130 strength 31–34, 37, 38 faunal evaluation 125 results discussed 36–43 cladistic analysis 131–133, 133–136 Dimorphodon spp. 30, 31, 32, 36, 42, 46, 52 comparative anatomy 136–138 D. macronyx 2, 27, 32, 35, 36, 39, 40, 41 geometric morphometrics 133, 136 Dinosaur Park Formation (Canada) 14,16 summary overview 142–144 dinosaurs, as pterosaur predators 15 systematic palaeontology 138–141 Diopecephalus spp. 125, 135 taxonomic review 126–129 comparative anatomy 136–137 stratigraphic review 130–131 D. kochi 129, 130, 130, 131, 141 D. rhamphastinus 140 gait and stance 45–46 systematics 138–140 gastric pellets 16 taxonomic review 126–128 gastroliths 10 Dorygnathus spp. 31, 119, 120, 136, 176 geometric morphometrics 133, 136, 139 D. banthensis 27, 32, 34, 35, 36, 39, 40, 41 Germanodactylus spp. Dsungaripteroidea 107, 149 G. cristatus 128, 129, 130, 130, 140, 141, 149 definition 154 G. kochi 141 taxonomy 154–157 G. rhamphastinus 27, 33, 36, 40, 41, 125, 128–129, Dsungaripterus spp. 35, 38, 43 130, 130, 140, 141 D. weii 27, 33, 36, 39, 40, 41 Germany 12, 14, 16, 95, 149 Gnathosaurinae 223 eggs Gnathosaurus spp. 31, 34, 37, 42 first found as fossil 1, 83–84 G. subulatus 27, 33, 36, 39, 40, 41, 130, 130 shell studies 83 Grinstead Clay Formation 223 embryos gut content analysis 8–10 discoveries 83 with pterosaur as prey 16 Largacito Formation specimen 85, 86, 87, 88 comparison with other sites 92 Haas, Jean Otto 159–160, 167 maturity criteria 91–92 Haenamichnus uhangriensis 11,12 measurements 87 Hamipterus 84 preservation 84–86 Haopterus 119 skeletal elements Harpactognathus gentryii 105, 120 INDEX 235

Hastings Group 223 Morrison Formation 105 Hatzegopteryx spp. 167 classification of finds 107–111 H. thambena 161 discussion of results 111 Hongshanopterus 119 crests 116–118 Howman, Rev. G.E. 2 dentition 116 humerus studies fangs 116 embryo 87 femur 111–114 Monofenestrata 107, 109, 115–116 humerus 115–116 Rhamphorhynchus 176, 176, 178, 181, 182 wing phalanx 116 Vesperopterylus 100, 100 fieldwork 105–106 summary of comparisons 119–120 ilia 173 Oxford Clay Formation 178 insects in pterosaur diets 13 Phu Kradung Formation 181–184 Istodactylus spp. 34 Solnhofen Formation 8, 10, 13, 14, 16, 149, 184 I. latidens 27, 33, 36, 40, 41, 119, 221, 223 I. sinensis 119 Kansas, Pteranodon localities 69–70 Italy 209–210, 211 Kepodactylus insperatus 105 wing metacarpals study Kimmeridge Clay Formation 175, 184 description 212–215 Korea 12 plate tectonic setting 212 Kunpengopterus sinensis 84 problems of assignment 215–217 stratigraphic setting 210–212 Lacusovagus magnificens 195, 203–205 Lagarcito Formation (Argentina) 10 Jagua Formation (Cuba) 184 embryo discoveries 83, 84, 85, 86, 87, 88 Javelina Formation (USA) 14 comparison with other sites 92 jaw bone maturity criteria 91–92 detailed descriptions measurements 87 UWGM2424 71 preservation 84–86 YPM VP2432 71–73, 73 skeletal elements YPM VP2525 74–75, 75, 76, 77–78, 77 limbs 89–90 YPM VP2738 73–74, 74 pectoral girdle 89 observations discussed 78–80 skull 86–87 jaw features, Disungariptids 149 vertebral column 87–89 jaw fragment 224, 225, 226 taxonomic assignment 90–91 jaw marks 11–12 Laopteryx 105 Jehol Group (China) 195 Las Hoyas Lagersta¨tte 13 Jeholopterus spp. 95, 96, 103 Liaoxipterus 119 Jiufotang Formation (China) 13, 95 limb descriptions 89–90 Jordan 159 locomotion, debates 45, 53–54 Er Ruseifa old phosphate mines 160, 161 Lonchodectes spp. Junggar Basin 149 L. compressirostris 229, 231 Jurassic specimens 2, 11, 12, 13, 16 L. sagittirostris 221, 227, 229 Jurassic Longchodectidae 221, 229 Franconia laminated limestone lagersta¨tten 125, 130 Longchodraco spp. 224–225, 231 faunal evaluation 125 Longchognathosaurus 149, 156, 157 cladistic analysis 131–133, 133–136 Ludodactylus sibbicki 9,13 comparative anatomy 136–138 geometric morphometrics 133, 136 Mancos Shale Formation (Utah) 187 summary overview 142–144 marine environment, pterosaur preservation 178 systematic palaeontology 138–141 Marsh, O.C. 1 taxonomic review 126–129 mass of body, role in predation 19 stratigraphic review 130–131 membrane, wing see wing properties Lagarcito Formation 10 Mesadactylus ornithosphyos 105 embryo discoveries 83, 84, 85, 86, 87, 88 metacarpals comparison with other sites 92 China 100 maturity criteria 91–92 embryo 87 measurements 87 Italy 212–215 preservation 84–86 Jordan 161, 161, 163–164 skeletal elements USA 188–189, 188, 189, 190–191 limbs 89–90 metatarsals 101 pectoral girdle 89 Mexico, Cerro del Pueblo Formation 11,12 skull 86–87 modelling see skull modelling vertebral column 87–89 Mongolia 95, 149 taxonomic assignment 90–91 Monofenestrata 107, 130, 133, 135, 136, 144 236 INDEX

Montanazhdarcho minor 216 O. microdon 229 Mooreville Chalk Formation 15 O. nobilis 221 Mo¨rnsheim Limestone Formation 141 O. oweni 229 morphometrics, geometric 133, 136 O. sagittirostris 229 Morrison Formation (USA) 105 O. scaphorhynchus 229 classification of finds 107–111 O. tenuirostris 229 discussion of results 111 Orthniocheiroidea 223 crests 116–118 Owen, Richard 1 dentition 116 Oxford Clay Formation 171, 171–172 fangs 116 stratigraphy 172 femur 111–114 humerus 115–116 palaeoecology 20 wing phalanx 116 Palaeornis cliftii 221, 223, 227 fieldwork 105–106 patagium 57 summary of comparisons 119–120 peck marks 12 musculature reconstruction pectoral girdle 89 Vectidraco daisymorrisae 46 pelvic girdle 101 methods 47 pelvis, musculature 45–46 results 47–51 perching behaviour 95, 101 results discussed 51–54 pes prints 11,12 Musculus adductores (1 and 2) 49, 50, 51, 53 Petrodactylus rhamphastinus 140 Musculus ambiens 49, 50, 51, 53 phalanx 87, 177, 177–178, 177, 224, 225, 227 Musculus caudofemoralis 49, 51, 52 Phobetor 155 Musculus flexor tibialis externus 48, 49, 50,52 Phosphatodraco mauritanicus 168 Musculus flexor tibialis internus 48, 49, 50,52 Phu Kradung Formation (Thailand) 181 Musculus iliofemoralis 50,51 phylogeny 18, 184, 199–201, 200, 201–205 Musculus iliofibularis 48, 49, 50,52 Pierre Shale Formation 17 Musculus iliotibialis 47–48, 49, 50,51 pneumatic spaces 71 Musculus ischiotrochantericus 51 Posidonienschiefer 184 Musculus puboischiofemoralis externus 49, 50, 51, 52, 53 predation Musculus puboischiofemoralis internus 48, 49, 50, 52, 53 on pterosaurs 15–17 possible pterosaur behaviour 13 narial openings 25 v. scavenging 19 Nemicolopterus crypticus 13 Preondactylus spp. 25, 42 Neoazhdarchia 223 P. buffaninii 27, 32, 34, 35, 36, 39, 40, 41 Nesodactylus spp. 176 prepubis 126, 131, 133, 136, 137, 138, 139, 140, Niobrara Formation (USA) 10, 14,15 141, 142, 143 Pteranodon descriptions 69–70 prod marks 11–12 jaw bone studies propatagium 57, 58 detailed descriptions Pteraichnus spp. 11, 12, 45 UWGM2424 71 Pteranodon spp. 9, 10, 15, 45, 216–217, 228 YPM VP2432 71–73, 73 P. ingens 27, 33, 36, 39, 40, 41 YPM VP2525 74–75, 75, 76, 77–78, 77 P. longiceps 70, 70 YPM VP2738 73–74, 74 P. sternbergi 70 observations discussed 78–80 skull strength 31, 32, 37, 42 Noripterus complicidens 119 wingspan 69 identification 150–151, 150, 151, 152 pteranodontoids 189–190, 191 systematics 153 Pterodactyloid pterosaur 229 taxonomy 154–157 Pterodactylus spp. 30, 42, 125, 176 Nusplingen limestone 14,16 P. antiquus 27, 33, 36, 39, 40, 41, 125, 130, 130 Nyctosaurus spp. P. cristatus 141 N. lamegoi 216 P. eurychirus 130 N. gracilis 213 P. kochi 125, 126–128, 141 P. longicollum 128 Oolithes bathonicae 83 P. micronyx 128 Oolithes sphaericus 83 P. montanus 105 orbit modelling 30 P. propinquus 12 Ornithocephalus ramphastinus 140 P. rhamphastinus 140 Ornithocheirus spp. 229 P. rhamphatilus 140 O. compressirostris 229 P. sagittirostris 229 O. curtus 221 P. scolopaciceps 125 O. daviesii 229 P. vulturinus 130 O. giganteus 229 Pterodaustro spp. 25, 31, 34, 42 O. machaeorhynchus 229 P. guinazui 10, 27, 33, 36, 40, 41,84 INDEX 237 pteroid 58, 100 orbit 35–36, 40 Pterorhynchus 184 stiffness 31 Pterosauria 107, 108 strength 31–34, 37, 38 cladistics 131–133, 133, 134, 135–136 results discussed 36–43 Purbeck Limestone Formation 11 Solnhofen Formation 8, 10, 13, 14, 16, 149, 184 Purbeckopus pentadactylus 11,12 South Dakota 17 Spain 13 Qinglongopterus spp. 184 stance and gait 45–46 quadrate inclination 37, 42 stiffness, wing membrane 59 quadrupedal stance 45 stomach contents analysis 10 Quetzalcoatlus spp. 16, 31, 36, 47, 83 subtree pruning and regrafting (SPR) distance Q. northropi 27, 33, 36, 40, 41, 161 132, 136, 137 Summerville Formation 11 radius 87, 100, 164, 166, 188, 188 sweeping marks 12 Raeticodactylus 25, 38 symposia 3 regurgitates 10, 13 syncarpal 227 with pterosaur as prey 16 synsacrals 173 rhamphorhynchines 184–185 systematics 8 Rhamphorhynchus spp. 13, 31, 36, 37, 120, 175 Arambourgiania philadelphiae 160–161 R. jessoni 171, 172–175, 173 Diopecephalus 138–140 R. kokeni 140 Lonchodraco 224–225 R. muensteri 8, 9, 10, 16, 18, 26, 27, 32, 34, 35, 36, Noripterus 153 40, 41 PRC64 181–184 Robinson–Foulds distances 132, 135–136, 136 Romania 14,16 tail drags 12 Romualdo Formation 195 tanglegrams 133, 135, 138 rostrum studies Tapejara wellnerhoferi 27, 33, 36, 38, 40, 41 detailed descriptions tapejarids 201 UWGM2424 71 teeth YPM VP2432 71–73, 73 Disungariptids 149 YPM VP2525 74–75, 75, 76, 77–78, 77 embedded in pterosaurs 15–16 YPM VP2738 73–74, 74 prey handling 25 observations discussed 78–80 Pterosauria (indet.), Morrison Formation 108–111, 110, 111, 116–118, 117, 120 St Leonards on Sea 229 Thailand specimen 181 Santana Formation 14, 16, 36 Tethys Ocean 210 Santana Group 195 Thailand, Jurassic pterosaur 181–184 Santanadactylus spp. Tiaojishan Formation 83 S. araripenis 228 tibia 87, 101, 174 S. pricei 228 toe, reversed 95, 101 S. spixi 228 Toolebuc Formation 14,16 Sao Khua Formation (Thailand) 181 tooth marks on pterosaurs 15–16 Scaphognathus spp. 30, 31, 120, 136 trace fossils 11–12 S. crassirostris 9, 27, 32, 34, 36, 40, 41 trackways 11 scapulocoracoid 97, 175–176, 175 Triassic, oldest pterosaur 8, 17, 69 scavenging behaviour, v. predation 19 Tropeognathus spp. 26 Sebes¸ Formation (Romania) 14,16 T. mesembrinus 27, 33, 36, 40, 41 Seeley, Harry Govier 1, 2 Tunbridge Wells Sand Formation 221, 223 Sericipterus spp. 176, 184 Serradraco spp. 231 ulna S. sagittirostris 223, 229, 230, 231 embryo measurements 87 sharks, as predators on pterosaurs 15–16, 19 Ferron Sandstone 187–188, 188, 190 skull Oxford Clay Formation 176, 176 bending moments 25 Upper Tunbridge Wells Sand Formation 224, bone pneumatization 69 225, 227 description 86–87 Vesperopterylus lamadongensis 100, 100 stresses on 26 Utah 11, 187 see also jaw bone Utahdactylus kateae 105 skull modelling methods Vectidraco daisymorrisae 46 geometry 26–27 muscle study orbital area 30 methods 47 stiffness 27–30 results 47–51 results 30–31 results discussed 51–54 238 INDEX

Vesperopterylus lamadongensis 95–96 Wight, Isle of 221 comparative anatomy 101, 103 wing metacarpals description 96, 100–101 new Italian specimen measurements 100 described 212–215 palaeoenvironment 101 problems of assignment 215–217 phylogeny 102, 103 wing phalanx 177, 177–178, 177 skeleton 96, 98, 99 wing properties 57, 89, 100 vertebrae and vertebral column 224, 225 bone morphology 57–58 Bexhill specimen 225–227 flutter limits 63–64 caudal and dorsal 173 maximum loading 61–63 cervical vertebra 162, 164–165, 196–199, membrane strain and deformation 197, 198, 199 64–66, 66 cranial fragment 165, 166–167 membrane tension 61, 62, 63, 65, 66 description 87–89, 96 morphology 58 indeterminate 174 section analysis 58–59, 59 thoracic or cervical 166, 167 structural model 60–61 Villaggio del Pescatore 209, 210 wingspan 161 Volgadraco bogolubovi 160 estimates 70, 84, 217 wrist 58 Wadhurst Clay Formation 221, 223 wukongopterids 120 Weald Clay Group 223 Wealden Supergroup 221, 223 Young’s Modulus 59 Wessex Formation 221, 223 Whitby Mudstone Formation 184 Zorzino Limestone Formation 8