Anhangueria (or anhanguerians) is a group of pterosaurs belonging to the clade Pteranodontoidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stages), around 140 to 92.5 million years ago. Anhangueria was named by paleontologists Taissa Rodrigues and Alexander Kellner in a review of Ornithocheirus species in 2013, they defined the clade as a branch-based taxon consisting of all pteranodontoids more closely related to Anhanguera blittersdorffi than to Istiodactylus latidens and Cimoliopterus cuvieri.[1]

Anhanguerians
Temporal range: Valanginian-Turonian,
~140–92.5 Ma
Iberodactylus and Hamipterus snouts.jpg
Snouts of the hamipterids Iberodactylus (left) and Hamipterus (right)
Guidraco venator 324.jpg
Holotype skull of the anhanguerid Guidraco
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Suborder: Pterodactyloidea
Clade: Ornithocheirae
Clade: Anhangueria
Rodrigues & Kellner, 2013
Subgroups

ClassificationEdit

Anhangueria originally only contained the genera Brasileodactylus, Camposipterus, Cearadactylus, Ludodactylus as well as the family Anhangueridae,[1] however, recent analyses had recovered the family Hamipteridae within this clade as well.[2][3][4][5][6]

In 2014, paleontologist Brian Andres and colleagues assigned more groups and genera within this clade, this included Guidraco, the subfamily Boreopterinae, and the clade Ornithocheirae, which was further divided into the families Anhangueridae and Ornithocheiridae.[7] This topology was later followed by a few other studies.[8][9] The cladogram of the phylogenetic analysis by Andres and colleagues is shown below:

Lanceodontia

Istiodactylidae

Anhangueria

Guidraco

Boreopterinae

Boreopterus

Zhenyuanopterus

Cearadactylus

Brasileodactylus

Ludodactylus

Ornithocheirae
Anhangueridae

Liaoningopterus

Anhanguera

Ornithocheiridae

Tropeognathus

Ornithocheirinae

Coloborhynchus

Ornithocheirus

Later, in 2019, a different phylogenetic analysis, this time conducted by Borja Holgado and colleagues, focused on derived pterodactyloids with particular emphasis on anhanguerians. In this analysis, Anhangueria consisted of Camposipterus and the families Anhangueridae and Hamipteridae, the clade Ornithocheirae was recovered as a more inclusive group consisting of Ornithocheirus and Cimoliopterus as basal members, as well as the Anhangueria in the most derived position. This classification is supported by the expansion of the premaxillary tip (a spoon-like expansion at the end of the snout) with a high jaw end.[2] Many analyses afterwards have followed this concept,[3][5][10] although some had recovered Camposipterus outside the Anhangueria (within the clade Targaryendraconia), but still within the Ornithocheirae.[4][6] The cladogram of the analysis by Holgado and colleagues in shown below:

 
Skull comparisons of Hamipterus (A and B), Ludodactylus (C), Caulkicephalus (D), Tropeognathus (E and F), Anhanguera (G and H), and Uktenadactylus (I and J)
Ornithocheirae

Ornithocheirus

Cimoliopterus

Anhangueria

Camposipterus

Hamipteridae

Hamipterus

Iberodactylus

Anhangueridae

Tropeognathus

Coloborhynchinae

Coloborhynchus

Uktenadactylus

Siroccopteryx

Anhanguerinae

Caulkicephalus

Guidraco

Ludodactylus

Liaoningopterus

Cearadactylus

Maaradactylus

Anhanguera

ReferencesEdit

  1. ^ a b Rodrigues, T.; Kellner, A. (2013). "Taxonomic review of the Ornithocheirus complex (Pterosauria) from the Cretaceous of England". ZooKeys (308): 1–112. doi:10.3897/zookeys.308.5559. PMC 3689139. PMID 23794925.
  2. ^ a b 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
  3. ^ a b 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.
  4. ^ a b Rodrigo V. Pêgas, Borja Holgado & Maria Eduarda C. Leal (2019) On Targaryendraco wiedenrothi gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids, Historical Biology, doi:10.1080/08912963.2019.1690482
  5. ^ a b 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). doi:10.1038/s41598-019-54042-z.
  6. ^ a b Holgado, B.; Pêgas, R.V. (2020). "A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae". Acta Palaeontologica Polonica. 65. doi:10.4202/app.00751.2020.
  7. ^ 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.
  8. ^ 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
  9. ^ Pentland, Adele H.; Poropat, Stephen F.; Tischler, Travis R.; Sloan, Trish; Elliott, Robert A.; Elliott, Harry A.; Elliott, Judy A.; Elliott, David A. (December 2019). "Ferrodraco lentoni gen. et sp. nov., a new ornithocheirid pterosaur from the Winton Formation (Cenomanian–lower Turonian) of Queensland, Australia". Scientific Reports. 9 (1): 13454. doi:10.1038/s41598-019-49789-4. ISSN 2045-2322. PMC 6776501. PMID 31582757.
  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.