Anhangueria
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Anhangueria (or anhanguerians) is a group of pterosaurs belonging to the clade Pteranodontoidea. Fossil remains of this group date back from the
Early Early may refer to: History * The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.: ** Early Christianity ** Early modern Europe Places in the United States * Early, Iowa * Early, Texas * Early ...
to
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
periods (
Valanginian In the geologic timescale, the Valanginian is an age or stage of the Early or Lower Cretaceous. It spans between 139.8 ± 3.0 Ma and 132.9 ± 2.0 Ma (million years ago). The Valanginian Stage succeeds the Berriasian Stage of the Lower Cretace ...
to
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded b ...
stages), around 140 to 92.5 million years ago. Anhangueria was named by paleontologists Taissa Rodrigues and
Alexander Kellner Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...
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 ''Anhanguera'' () is a genus of pterodactyloid pterosaur known from the Early Cretaceous (Albian age, 125 to 112 million years ago) Romualdo Formation of Brazil and the Late Cretaceous ( Cenomanian age, 98 to 92.5 million years ago) Kem Kem G ...
'' than to ''
Istiodactylus latidens ''Istiodactylus'' is a genus of pterosaur that lived during the Early Cretaceous period, about 120 million years ago. The first fossil was discovered on the English Isle of Wight in 1887, and in 1901 became the holotype specimen of a new species ...
'' and '' Cimoliopterus cuvieri''.


Classification

Anhangueria originally only contained the
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
''
Brasileodactylus ''Brasileodactylus'' a genus of pterosaur from the Aptian-age (Early Cretaceous period) lower Santana formation of Chapada do Araripe, Ceará, Brazil. The genus was named by paleontologist Alexander Wilhelm Armin Kellner in 1984. The genus nam ...
'', '' Camposipterus'', '' Cearadactylus'', ''
Ludodactylus ''Ludodactylus'' (meaning "play finger") is a genus of anhanguerid pterodactyloid pterosaur from the Early Cretaceous period (Aptian stage) of what is now the Crato Formation of the Araripe Basin in Ceará, Brazil. The type and only species i ...
'' as well as the family Anhangueridae, however, recent analyses had recovered the family Hamipteridae within this clade as well. In 2014, paleontologist Brian Andres and colleagues assigned more groups and genera within this clade, this included ''
Guidraco ''Guidraco'' (Chin. ''gui'' (鬼) "malicious ghost" + Lat. ''draco'' "dragon") is an extinct genus of toothed pterodactyloid pterosaur known from the Early Cretaceous of Liaoning Province, northeast China. According to many recent studies, ''Gu ...
'', the subfamily
Boreopterinae Boreopteridae (meaning "northern wings") is a group of pterodactyloid pterosaurs from the Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China. Classification In 2006, Lü Junchang and colleagues named the clade Boreopteridae for t ...
, and the clade
Ornithocheirae Ornithocheirae is an extinct clade of pteranodontoid pterosaurs from the Early Cretaceous to the Late Cretaceous (Valanginian to Turonian stages) of Asia, Europe, North America and South America. It was named by Harry Seeley in 1870 as a famil ...
, which was further divided into the families Anhangueridae and
Ornithocheiridae Ornithocheiridae (or ornithocheirids, meaning "bird hands") is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the Early to Late Cr ...
. This topology was later followed by a few other studies.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.
The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
of the
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
by Andres and colleagues is shown below: Later, in 2019, a different phylogenetic analysis, this time conducted by Borja Holgado and colleagues, focused on derived
pterodactyloid Pterodactyloidea (derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger" meaning "winged finger", "wing-finger" or "finger-wing") is one of the two traditional ...
s 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 ''Cimoliopterus'' is a genus of pterosaur that lived during the Late Cretaceous in what is now England and the United States. The first known specimen, consisting of the front part of a snout including part of a crest, was discovered in the ...
'' as basal members, as well as the Anhangueria in the most derived position. This classification is supported by the expansion of the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
ry tip (a spoon-like expansion at the end of the snout) with a high jaw end.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 Many analyses afterwards have followed this concept,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). . although some had recovered ''Camposipterus'' outside the Anhangueria (within the clade
Targaryendraconia Targaryendraconia is an extinct clade of lanceodontian pterosaurs that lived from the Early to Late Cretaceous period in Europe, North America, South America, and Australia. Classification Below is a cladogram following a topology by Pêgas ...
), but still within the Ornithocheirae.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, The cladogram of the analysis by Holgado and colleagues in shown below:


References

{{Taxonbar, from=Q21008914 Pteranodontoids Early Cretaceous first appearances Late Cretaceous extinctions