Ornithocheiroidea

Ornithocheiroidea (or ornithocheiroids) is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods (Valanginian to Maastrichtian stages), with fossil remains found all over the world except Antarctica.

Ornithocheiroids
Temporal range: Early - Late Cretaceous,
~140–66 Ma Possible Late Jurassic record[1]
Thalassodromeus in Japan.jpg
Restored skeleton of Thalassodromeus sethi, with Anhanguera behind
Tropeognathus mesembrinus MN 01.jpg
Skeletal cast of Tropeognathus mesembrinus in the National Museum of Brazil
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Suborder: Pterodactyloidea
Infraorder: Eupterodactyloidea
Clade: Ornithocheiroidea
Seeley, 1870
Subgroups

Ornithocheiroids were the most advanced group of pterosaurs, as the group includes the clade Azhdarchoidea, of which its members lived until the Maastrichtian stage of the Late Cretaceous, around 66 million years ago. Notable pterosaurs from this group include the pteranodontians Pteranodon and Nyctosaurus, the ornithocheirid Ornithocheirus, the anhanguerid Tropeognathus, as well as the azhdarchids Hatzegopteryx and Quetzalcoatlus.

ClassificationEdit

The name Ornithocheiroidea was originally defined as an apomorphy-based taxon by Christopher Bennett in 1994. It was given a relationship-based definition in 2003 by Alexander Kellner, who defined it as the least inclusive clade containing Anhanguera blittersdorffi, Pteranodon longiceps, Dsungaripterus weii, and Quetzalcoatlus northropi.[2] Later that year, David Unwin suggested a more restrictive definition, in which the clade only contains Pteranodon longiceps, Istiodactylus latidens, and their descentants.[3] Brian Andres (2008, 2010, 2014) in his analyses, defined Ornithocheiroidea using the definition of Kellner (2003) to avoid confusion with similarly-defined groups, like Pteranodontoidea.[4][5]

Below is a cladogram showing the results of a phylogenetic analysis presented by Longrich and colleagues in 2018. They found Ornithocheiroidea to consist of the clades Pteranodontoidea and Azhdarchoidea, as well as the genus Piksi.[6]

Eupterodactyloidea

Haopterus gracilis

Ornithocheiroidea

Piksi barbarulna

Pteranodontoidea
Pteranodontia
Pteranodontidae

Tethydraco regalis

Pteranodon longiceps 

Pteranodon sternbergi

Nyctosauridae

Alamodactylus byrdi

Volgadraco bogolubovi

Cretornis hlavaci

Alcione elainus

Simurghia robusta

Muzquizopteryx coahuilensis

Barbaridactylus grandis

Nyctosaurus lamegoi

Nyctosaurus nanus

Nyctosaurus gracilis 

Ornithocheiromorpha

Hongshanopterus lacustris

Lonchodectidae

Lonchodraco giganteus

Lonchodectes compressirostris

Boreopteridae

Boreopterus cuiae

Zhenyuanopterus longirostris

Lanceodontia
Istiodactylidae

Nurhachius ignaciobritoi

Liaoxipterus brachyognathus

Istiodactylus sinensis

Istiodactylus latidens

Aetodactylus halli

Cimoliopterus dunni

Cimoliopterus cuvieri

Anhangueria

Guidraco

Ludodactylus 

Cearadactylus

Brasileodactylus

Ornithocheirae

Anhangueridae 

Ornithocheiridae

Azhdarchoidea
Tapejaromorpha

Bennettazhia oregonensis

Tapejaridae
Tapejarinae

Tapejara wellnhoferi

Europejara olcadesorum

Vectidraco daisymorrisae

Caiuajara dobruskii

Tupandactylus navigans

Tupandactylus imperator 

Bakonydraco galaczi 

"Huaxiapterus" benxiensis

"Huaxiapterus" corollatus

Eopteranodon lii

Huaxiapterus jii

Sinopterus dongi

Neoazhdarchia
Dsungaripteromorpha
Dsungaripteridae
Dsungaripterinae

Dsungaripterus weii

Domeykodactylus ceciliae 

Noripterinae

Noripterus parvus

Noripterus complicidens

Thalassodromidae

Tupuxuara longicristatus 

Tupuxuara leonardii

Thalassodromeus sethi

Alanqa saharica

Aerotitan sudamericanus 

Neopterodactyloidea
Chaoyangopteridae

Microtuban altivolans

Chaoyangopterinae

Shenzhoupterus

Chaoyangopterus

Jidapterus

Eoazhdarcho liaoxiensis

Radiodactylus langstoni

Azhdarchidae

Montanazhdarcho minor

Azhdarcho lancicollis

Phosphatodraco mauritanicus 

Aralazhdarcho bostobensis

Eurazhdarcho langendorfensis

aff. Quetzalcoatlus

Zhejiangopterus linhaiensis

Arambourgiania philadelphiae

Hatzegopteryx thambema

Quetzalcoatlus spp. 

In 2019, a phylogenetic analysis conducted by Kellner and colleagues had recovered Ornithocheiroidea as the sister taxon of the Archaeopterodactyloidea, and consisting of the clades Tapejaroidea and Pteranodontoidea.[7] Several recent studies have followed this or a similar concept.[8][9][10] The cladogram of the analysis by Kellner and colleagues is presented below:

Pterodactyloidea

Archaeopterodactyloidea 

Ornithocheiroidea
Pteranodontoidea
Pteranodontia
Pteranodontidae

Pteranodon longiceps 

Tethydraco regalis

Nyctosauridae

Nyctosaurus gracilis 

Muzquizopteryx coahuilensis

Lanceodontia

Hongshanopterus lacustris

Istiodactylidae

Nurhachius ignaciobritoi

Istiodactylinae

Istiodactylus latidens

Istiodactylus sinensis

Haopterus gracilis

Lonchodraco giganteus

Ikrandraco avatar

Ornithocheirae

Ornithocheirus simus 

Cimoliopterus cuvieri

Anhangueria

Camposipterus sedgwickii

Hamipteridae

Hamipterus tianshanensis

Iberodactylus andreui

Anhangueridae

Tropeognathus mesembrinus

Coloborhynchinae

Coloborhynchus clavirostris

Uktenadactylus wadleighi

Anhanguerinae

Caulkicephalus trimicrodon

Guidraco venator

Ludodactylus sibbicki 

Liaoningopterus gui 

Cearadactylus atrox

Maaradactylus kellneri

AMNH 22555

Anhanguera blittersdorffi

Anhanguera piscator

Tapejaroidea
Dsungaripteridae

Dsungaripterus weii

Noripterus parvus

Azhdarchoidea
Azhdarchidae

Azhdarcho lancicollis

Quetzalcoatlus sp. 

Zhejiangopterus linhaiensis

Chaoyangopteridae

Chaoyangopterus zhangi

Jidapterus edentus

Shenzhoupterus chaoyangensis

Tapejaromorpha

Keresdrakon vilsoni

Tapejaridae
Thalassodrominae

Thalassodromeus sethi

Tupuxuara leonardii

Tapejarinae

Caupedactylus ybaka

Aymberedactylus cearensis

Eopteranodon lii

"Huaxiapterus" benxiensis

"Huaxiapterus" corollatus

Sinopterus dongi

Tapejarini

Europejara olcadesorum

Caiuajara dobruskii

Tapejara wellnhoferi

Tupandactylus imperator 

ReferencesEdit

  1. ^ Unwin, David M.; Heinrich, Wolf-Dieter (1999). "On a pterosaur jaw from the Upper Jurassic of Tendaguru (Tanzania)". Mitteilungen aus dem Museum für Naturkunde in Berlin, Geowissenschaftliche Reihe. 2: 121–134.
  2. ^ Kellner, A. W. A., (2003): Pterosaur phylogeny and comments on the evolutionary history of the group. pp. 105-137. — in Buffetaut, E. & Mazin, J.-M., (eds.): Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347
  3. ^ Unwin, D. M., (2003): On the phylogeny and evolutionary history of pterosaurs. pp. 139-190. — in Buffetaut, E. & Mazin, J.-M., (eds.): Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347 doi:10.1144/GSL.SP.2003.217.01.11
  4. ^ Andres, Brian Blake (2014). Systematics of the Pterosauria. Yale University. p. 366. A preview that shows the cladogram without clade names
  5. ^ Andres, B.; Clark, J.; Xu, X. (2014). "The Earliest Pterodactyloid and the Origin of the Group". Current Biology. 24 (9): 1011–6. doi:10.1016/j.cub.2014.03.030. PMID 24768054.
  6. ^ Longrich, N.R., Martill, D.M., and Andres, B. (2018). Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary. PLoS Biology, 16(3): e2001663. doi:10.1371/journal.pbio.2001663
  7. ^ Kellner, Alexander W. A.; Weinschütz, Luiz C.; Holgado, Borja; Bantim, Renan A. M.; Sayão, Juliana M. (August 19, 2019). "A new toothless pterosaur (Pterodactyloidea) from Southern Brazil with insights into the paleoecology of a Cretaceous desert". Anais da Academia Brasileira de Ciências. 91 (suppl 2): e20190768. doi:10.1590/0001-3765201920190768. ISSN 0001-3765. PMID 31432888.
  8. ^ Kellner, Alexander W. A.; Caldwell, Michael W.; Holgado, Borja; Vecchia, Fabio M. Dalla; Nohra, Roy; Sayão, Juliana M.; Currie, Philip J. (2019). "First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity". Scientific Reports. 9 (1): 17875. Bibcode:2019NatSR...917875K. doi:10.1038/s41598-019-54042-z. PMC 6884559. PMID 31784545.
  9. ^ Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", Scientific Reports 9: 4940 doi:10.1038/s41598-019-41280-4
  10. ^ Jiang, Shun-Xing; Zhang, Xin-Jun; Cheng, Xin; Wang, Xiao-Lin (2020). "A new pteranodontoid pterosaur forelimb from the upper Yixian Formation, with a revision of Yixianopterus jingangshanensis" (PDF). Vertebrata PalAsiatica. doi:10.19615/j.cnki.1000-3118.201124.