Tapejaridae
Encyclopedia
Tapejaridae are a family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 of pterodactyloid pterosaurs from the early Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

 period. Members are currently known from Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...

 and China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

, where the most primitive genera are found, indicating that the family has an Asian origin.

Description

Tapejarids were small to medium-sized pterosaurs with several unique, shared characteristics, mainly relating to the skull. Most tapejarids possessed a bony crest arising from the snout (formed mostly by the premaxillary bones of the upper jaw tip). In some species, this bony crest is known to have supported an even larger crest of softer, fibrous tissue that extends back along the skull. Tapejarids are also characterized by their large nasoantorbital fenestra, the main opening in the skull in front of the eyes, which spans at least half the length of the entire skull in this family. Their eye sockets were small and pear-shaped.

Taxonomy

Tapejaridae may be composed of two subfamilies: a Tapejarinae of "Huaxiapterus" corollatus
Huaxiapterus
"Huaxiapterus" is a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. It is the second genus of tapejarid from this formation, after Sinopterus, and may be more derived. Two species are known.The type species of...

, Sinopterus
Sinopterus
Sinopterus was a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. Three species have been classified in this genus, though only two are generally considered to be valid...

, Tapejara, Tupandactylus
Tupandactylus
Tupandactylus is a genus of tapejarid pterodactyloid pterosaur from the Early Cretaceous Crato Formation of Brazil. It is notable for its large cranial crest, composed partly of bone and partly of soft tissue...

, and a Thalassodrominae of Thalassodromeus
Thalassodromeus
Thalassodromeus was a large pterodactyloid pterosaur found in northeastern Brazil.The genus was named in 2002 by Alexander Kellner and Diogenes de Almeida Campos. The type species is Thalassodromeus sethi. The genus name is derived from Greek thalasse, "sea" and dromaios, "runner", in reference to...

and Tupuxuara
Tupuxuara
Tupuxuara is a genus of large, crested, toothless pterodactyloid pterosaur.The genus was in 1988 named and described by Alexander Kellner and Diogenes de Almeida Campos. The type species is Tupuxuara longicristatus. The genus name refers to a familiar spirit from the mythology of the Tupi...

. Some studies, such as one by Lü and colleagues in 2008, have found that the thalassodromines are more closely related to the azhdarchids
Azhdarchidae
Azhdarchidae is a family of pterosaurs known primarily from the late Cretaceous Period, though an isolated vertebrae apparently from an azhdarchid is known from the early Cretaceous as well...

 proper than to the tapejarids, and have placed them in their own family (which has sometimes been referred to as Tupuxuaridae, though Thalassodrominae was named first). At least one study has also found that the Chaoyangopteridae
Chaoyangopteridae
The Chaoyangopteridae are a family of pterosaurs within the Azhdarchoidea.The clade Chaoyangopteridae was first defined in 2008 by Lü Junchang and David Unwin as: "Chaoyangopterus, Shenzhoupterus, their most recent common ancestor and all taxa more closely related to this clade than to Tapejara,...

, often found to be closer to azhdarchids, represent a lineage within the Tapejaridae, more closely related to the tapejarines than to the thalassodromines. Felipe Pinheiro and colleagues (2011) reclassified the group as a subfamily of Tapejaridae, Chaoyangopterinae, for this reason.
  • Family Tapejaridae
    • Subfamily Chaoyangopterinae?
      • Chaoyangopterus
        Chaoyangopterus
        Chaoyangopterus is a genus of azhdarchoid pterodactyloid pterosaur known from a partial skeleton found in Liaoning, China. It was found in rocks of the Aptian-age Lower Cretaceous Jiufotang Formation of Dapingfang, Chaoyang....

      • Eoazhdarcho
        Eoazhdarcho
        Eoazhdarcho is a genus of azhdarchoid pterodactyloid pterosaur named in 2005 by Chinese paleontologists Lü Junchang and Ji Qiang. The type species is Eoazhdarcho liaoxiensis. The genus name combines a Greek eos, "dawn" with the name of the genus Azhdarcho, with the implication it was an early...

        ?
      • Eopteranodon
        Eopteranodon
        Eopteranodon is a genus of azhdarchoid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Yixian Formation of Beipaio City, Liaoning, China....

        ?
      • Jidapterus
        Jidapterus
        Jidapterus is a genus of azhdarchoid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China....

      • Shenzhoupterus
        Shenzhoupterus
        Shenzhoupterus is a genus of chaoyangopterid azhdarchoid pterodactyloid pterosaur from the Lower Cretaceous Jiufotang Formation of Liaoning, China....

    • Subfamily Tapejarinae
      • "Huaxiapterus
        Huaxiapterus
        "Huaxiapterus" is a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. It is the second genus of tapejarid from this formation, after Sinopterus, and may be more derived. Two species are known.The type species of...

        "
      • Sinopterus
        Sinopterus
        Sinopterus was a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. Three species have been classified in this genus, though only two are generally considered to be valid...

      • Tapejara
      • Tupandactylus
        Tupandactylus
        Tupandactylus is a genus of tapejarid pterodactyloid pterosaur from the Early Cretaceous Crato Formation of Brazil. It is notable for its large cranial crest, composed partly of bone and partly of soft tissue...

    • Subfamily Thalassodrominae?
      • Thalassodromeus
        Thalassodromeus
        Thalassodromeus was a large pterodactyloid pterosaur found in northeastern Brazil.The genus was named in 2002 by Alexander Kellner and Diogenes de Almeida Campos. The type species is Thalassodromeus sethi. The genus name is derived from Greek thalasse, "sea" and dromaios, "runner", in reference to...

      • Tupuxuara
        Tupuxuara
        Tupuxuara is a genus of large, crested, toothless pterodactyloid pterosaur.The genus was in 1988 named and described by Alexander Kellner and Diogenes de Almeida Campos. The type species is Tupuxuara longicristatus. The genus name refers to a familiar spirit from the mythology of the Tupi...


Phylogeny

The exact relationships of tapejarids to one another and to other azhdarchoid
Azhdarchoidea
Azhdarchoidea is a group of pterosaurs within the suborder Pterodactyloidea.-Classification:Listing of families and genera after Unwin 2006 except where specified.* Superfamily Azhdarchoidea** Genus "Aurorazhdarcho"** Family Chaoyangopteridae...

 pterosaurs has historically been unclear, with different studies producing significantly different cladogram
Cladogram
A cladogram is a diagram used in cladistics which shows ancestral relations between organisms, to represent the evolutionary tree of life. Although traditionally such cladograms were generated largely on the basis of morphological characters, DNA and RNA sequencing data and computational...

s (family trees). It is also unclear exactly which pterosaurs belong to the Tapejaridae; some researchers have found the thalassodromines and chaoyangopterines to be members of this family, while other studies have found them to be more closely related to the azhdarchids
Azhdarchidae
Azhdarchidae is a family of pterosaurs known primarily from the late Cretaceous Period, though an isolated vertebrae apparently from an azhdarchid is known from the early Cretaceous as well...

 (in the clade Neoazhdarchia). Some studies have even allowed the possibility that the "tapejarids" as traditionally thought of are paraphyletic, that is, they may not form a natural group but instead represent sequential branches of the tree leading to the more advanced neoazhdarchians. Below are two alternate cladograms: the first, presented by Felipe Pinheiro and colleagues in 2011, shows the tapejarids as a monophyletic clade including the thalassodromines and chaoyangopterines. The second, presented by Dave Martill and Darren Naish in 2006, shows them as a paraphyletic grade.
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