Tapejaridae

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Tapejaridae
Temporal range: Early to Late Cretaceous, 125–85  Ma
Tapejaridae skull comparison.png
Collection of various tapejarid skulls to scale with one another. From left to right, top to bottom:

Torukjara bandeirae , Caiuajara dobruskii , Tupandactylus imperator , Tapejara wellnhoferi , Huaxiadraco corollatus , and Sinopterus dongi

Contents

Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order:† Pterosauria
Suborder:† Pterodactyloidea
Clade:† Tapejaromorpha
Family:† Tapejaridae
Kellner, 1989
Type species
† Tapejara wellnhoferi
Kellner, 1989
Genera
Synonyms
  • Eutapejaria?
    (Pêgas et al., 2023)

Tapejaridae (from a Tupi word meaning "the lord of the ways") are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, [3] Spain, [4] the United States, [5] and China. The most primitive genera were found in China, indicating that the family has an Asian origin. [6]

Description

Reconstructed profiles of three Brazilian species; from top to bottom, Tapejara wellnhoferi (A), Tupandactylus navigans (B), and Tupandactylus imperator (C) Tapejarines mmartyniuk.png
Reconstructed profiles of three Brazilian species; from top to bottom, Tapejara wellnhoferi (A), Tupandactylus navigans (B), and Tupandactylus imperator (C)

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. [7] Studies of tapejarid brain cases show that they had extremely good vision, more so than in other pterosaur groups, and probably relied nearly exclusively on vision when hunting or interacting with other members of their species. [8] Tapejarids had unusually reduced shoulder girdles that would have been slung low on the torso, resulting in wings that protruded from near the belly rather than near the back, a "bottom decker" arrangement reminiscent of some planes. [8]

Biology

Tapejarids appear to have been arboreal, having more curved claws than other azhdarchoid pterosaurs and occurring more commonly in fossil sites with other arboreal flying vertebrates such as early birds. Tapejarids have long been speculated as having been frugivores or omnivores, based on their parrot-like beaks. [9] Direct evidence for plant-eating is known in a specimen of Sinopterus that preserves seeds in the abdominal cavity. The Barremian- Aptian distribution of some tapejarids may even be partially associated with the first radiation phase of the angiosperms, especially of the genus Klitzschophyllites which represents a more basal angiosperm. [10] [11]

Assuming thalassodromines are tapejarids, this clade also included raptorial, macropredatory species adapted to hunt proportionally large prey. [12] [13]

Classification

Tapejarids.jpg
Life restorations of various members of the Tapejaridae
Azhdarchoidea creta inf 120 maf brasile.jpg
Hind leg of an indeterminate tapejarid, with preserved foot pads, scales, and claw sheaths

Tapejaridae was defined by Alexander Kellner in 1989 as the clade containing both Tapejara and Tupuxuara, plus all descendants of their most recent common ancestor. As originally conceived, it was composed of two subfamilies: the Tapejarinae, consisting of Tapejara and its close relatives, and the Thalassodrominae, consisting of Thalassodromeus and Tupuxuara . [14]

Some studies, such as one by Lü and colleagues in 2008, have found that the thalassodromines are more closely related to the azhdarchids proper than to the tapejarids, [15] and have placed them in their own family (which has sometimes been referred to as Tupuxuaridae, [16] though Thalassodrominae was named first [14] ). At least one study has also found that the Chaoyangopteridae, 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. [7]

The exact relationships of tapejarids to one another and to other azhdarchoid pterosaurs has historically been unclear, with different studies producing significantly different cladograms (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, [7] [14] while other studies have found them to be more closely related to the azhdarchids (in the clade Neoazhdarchia). [17] Several studies have shown that the "tapejarids" as traditionally thought of (that is, including the classic examples of both Tapejara and Tupuxuara) are paraphyletic, and do not form a natural group, but instead represent sequential branches of the tree leading. In light of this discovery, several of the traditional names associated with the group have been re-defined. Martill and Naish proposed a revised definition for Tapejaridae, as all species more closely related to Tapejara than to Quetzalcoatlus . [17] Andres and colleagues did not follow this proposal, instead formally defining Tapejaridae as the clade Tapejara + Sinopterus. They also re-defined the Tapejarinae as all species closer to Tapejara than to Sinopterus, and added a new clade, Tapejarini, to include all descendants of the last common ancestor of Tapejara and Tupandactylus. [18] [19]

Below are two alternate cladograms: the first, presented by Andres and colleagues in 2014, found the a grouping of tapejarids at the base of the clade, with thalassodromines more closely related to azhdarchids, chaoyangopterids, and dsungaripterids, all of them within the group Neoazhdarchia. Their cladogram is shown on the left. [18] Later however, a number of studies had begun to favor Kellner's defition of the group, meaning that the members of the Thalassodrominae were reclassified within this group. [20] [21] An example of one of these studies is the one by Kellner and colleagues in 2019. The cladogram on the right shows their phylogenetic analysis. [22] In 2021 a new study focus on the most complete Tapejaridae, a skeleton Tupandactylus navigans, confirmed the 2019 topology [23] .

Related Research Articles

<i>Tapejara wellnhoferi</i> Genus of tapejarid pterosaur

Tapejara is a genus of Brazilian pterosaur from the Cretaceous Period. Tapejara crests consisted of a semicircular crest over the snout, and a bony prong which extended back behind the head. It was a small pterosaur, with a wingspan of approximately 1.23–1.3 metres (4.0–4.3 ft).

<i>Cearadactylus</i> Genus of anhanguerid pterosaur from the Early Cretaceous

Cearadactylus is a genus of large anhanguerid pterodactyloid pterosaur from the Romualdo Formation of Brazil, South America. Fossil remains of Cearadactylus dated back to the Albian stage of the Early Cretaceous period, about 112 million years ago. The only known species is C. atrox, described and named in 1985 by Giuseppe Leonardi and Guido Borgomanero. The name refers to the Brazilian state Ceará, and combines this with Greek daktylos, "finger", a reference to the wing finger of pterosaurs. The Latin atrox means "frightful", a reference to the fearsome dentition of the species.

<span class="mw-page-title-main">Azhdarchidae</span> Family of large azhdarchoid pterosaurs

Azhdarchidae is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cretaceous as well. Azhdarchids are mainly known for including some of the largest flying animals discovered, but smaller cat-size members have also been found. Originally considered a sub-family of Pteranodontidae, Nesov (1984) named the Azhdarchinae to include the pterosaurs Azhdarcho, Quetzalcoatlus, and Titanopteryx. They were among the last known surviving members of the pterosaurs, and were a rather successful group with a worldwide distribution. Previously it was thought that by the end of the Cretaceous, most pterosaur families except for the Azhdarchidae disappeared from the fossil record, but recent studies indicate a wealth of pterosaurian fauna, including pteranodontids, nyctosaurids, tapejarids and several indeterminate forms. In several analyses, some taxa such as Navajodactylus, Bakonydraco and Montanazhdarcho were moved from Azhdarchidae to other clades.

<i>Tupuxuara</i> Genus of azhdarchoid pterosaur from the Early Cretaceous

Tupuxuara is a genus of large, crested, and toothless pterodactyloid pterosaur from the Early Cretaceous period of what is now the Romualdo Formation of the Santana Group, Brazil, about 125 to 112 million years ago. Tupuxuara is a close relative of Thalassodromeus, and both form a group that is either called Thalassodrominae or Thalassodromidae.

<i>Thalassodromeus</i> Genus of azhdarchoid pterosaur from the Early Cretaceous

Thalassodromeus is a genus of pterosaur that lived in what is now Brazil during the Early Cretaceous period, about a hundred million years ago. The original skull, discovered in 1983 in the Araripe Basin of northeastern Brazil, was collected in several pieces. In 2002, the skull was made the holotype specimen of Thalassodromeus sethi by palaeontologists Alexander Kellner and Diogenes de Almeida Campos. The generic name means "sea runner", and the specific name refers to the Egyptian god Seth due to its crest being supposedly reminiscent of Seth's crown. Other scholars have pointed out that the crest was instead similar to the crown of Amon. A jaw tip was assigned to T. sethi in 2005, became the basis of the new genus Banguela in 2015, and assigned back to Thalassodromeus as the species T. oberlii in 2018, though other researchers consider it a valid genus. Another species was described in 2015 based on a supposed crest fragment, but this was later shown to be part of a turtle shell.

<i>Bakonydraco</i> Genus of tapejarid pterosaur

Bakonydraco is a genus of pterodactyloid pterosaur from the Late Cretaceous period of what is now the Csehbánya Formation of the Bakony Mountains, Iharkút, Veszprém, western Hungary.

<i>Eopteranodon</i> Genus of tapejarid pterosaur

Eopteranodon is a genus of tapejarid pterosaur from the Aptian-age Lower Cretaceous Yixian Formation of Beipiao City, Liaoning, China. The genus was named in 2005 by paleontologists Lü Junchang and Zhang Xingliao. The type species is Eopteranodon lii. A second species, E. yixianensis, was named in 2023, but its validity has been questioned.

<i>Chaoyangopterus</i> Genus of chaoyangopterid pterosaur from the Early Cretaceous

Chaoyangopterus is a genus of chaoyangopterid pterosaur known from a partial skeleton found in Liaoning, China. Chaoyangopterus was found in rocks dating back to the Aptian-age Lower Cretaceous Jiufotang Formation of Dapingfang, Chaoyang.

<i>Jidapterus</i> Genus of chaoyangopterid pterosaur from the Early Cretaceous

Jidapterus is a genus of chaoyangopterid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. The genus was in 2003 named by Dong Zhiming, Sun Yue-Wu and Wu Shao-Yuan. The type species is Jidapterus edentus. The genus name is derived from Jílín Dàxué or "Jilin University" and a Latinized Greek pteron, "wing". The specific name means "toothless" in Latin.

<span class="mw-page-title-main">Ornithocheiroidea</span> Clade of pterodactyloid pterosaurs

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

<span class="mw-page-title-main">Azhdarchoidea</span> Superfamily of ornithocheiroid pterosaurs

Azhdarchoidea is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, Tendaguripterus, that lived in the Late Jurassic period. The largest azhdarchoids include members of the family Azhdarchidae, examples of these are Quetzalcoatlus, Hatzegopteryx, and Arambourgiania. The Azhdarchoidea has been recovered as either closely related to the Ctenochasmatoidea, as the sister taxon of the Pteranodontoidea within the Ornithocheiroidea, or within the Tapejaroidea, which in turn was also within the Ornithocheiroidea.

<i>Tupandactylus</i> Genus of tapejarid pterosaur from the Early Cretaceous

Tupandactylus is a genus of tapejarid pterodactyloid pterosaur from the Early Cretaceous Crato Formation of Brazil.

<span class="mw-page-title-main">Chaoyangopteridae</span> Family of azhdarchoid pterosaurs

Chaoyangopteridae is a family of pterosaurs within the larger group Azhdarchoidea. Chaoyangopterids lived mostly during the Early Cretaceous period, though possible members, Microtuban, Xericeps and Argentinadraco, may extend the fossil range to the Late Cretaceous.

<span class="mw-page-title-main">Thalassodrominae</span> Subfamily of azhdarchoid pterosaurs from the Cretaceous period

Thalassodrominae or Thalassodromidae is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional members come from Brazil, however, other possible members also come from other places, including Morocco and Argentina. Thalassodrominae is considered either to be a subfamily within the pterosaur family Tapejaridae, or as a distinct family, Thalassodromidae, within the clade Neoazhdarchia, closely related to dsungaripterids or azhdarchids.

<i>Banguela</i> Genus of azhdarchoid pterosaur from the Early Cretaceous

Banguela is a genus of azhdarchoid pterosaur from the Early Cretaceous period of what is now Brazil. Only one species is known, Banguela oberlii.

<span class="mw-page-title-main">Tapejaroidea</span> Clade of ornithocheiroid pterosaurs

Tapejaroidea is a group of pterosaurs belonging to the clade Ornithocheiroidea. Tapejaroids lived from the Early to Late Cretaceous periods, with one possible member, Tendaguripterus, extending the fossil range to the Late Jurassic period. Tapejaroidea contains two groups, the Dsungaripteridae and the Azhdarchoidea, which in turn includes the azhdarchids, the group that contains some of the largest flying animals. The group was named by Brazilian paleontologist Alexander Wilhelm Armin Kellner in 1996.

<span class="mw-page-title-main">Tapejaromorpha</span> Clade of azhdarchoid pterosaurs

Tapejaromorpha is a group of pterosaurs within the clade Azhdarchoidea. The fossil remains of tapejaromorphs dated back to the Cretaceous period.

<i>Argentinadraco</i> Genus of azhdarchoid pterosaur from the Late Cretaceous

Argentinadraco is an extinct genus of azhdarchoid pterosaur from the Late Cretaceous Portezuelo Formation of Argentina. It contains a single species, A. barrealensis, named in 2017 by Alexander Kellner and Jorge Calvo. Argentinadraco is unusual for bearing a bottom jaw with a concave bottom edge, as well as a pair of ridges and depressions on the top surface. These features distinguish it from all other azhdarchoid groups, complicating its assignment, but it may belong to the Chaoyangopteridae. The ridges on the lower jaw may have been used to feed on small invertebrates in loose sediment within the system of lakes and rivers that it resided in.

<i>Wellnhopterus</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Wellnhopterus is an azhdarchid pterosaur recovered from the Late Cretaceous Javelina Formation in Texas that was previously identified as a thalassodromine. It consists of a set of upper and lower jaws, as well as some cervical vertebrae and a fragmentary long bone. In July 2021, the jaws were given the genus name "Javelinadactylus", with the type and only species as "J. sagebieli"; however, this article has now been retracted. In a paper published in December 2021, the complete holotype was independently named Wellnhopterus, with the only species being W. brevirostris. As of 2022, this is the formal name of this pterosaur.

<i>Huaxiadraco</i> Genus of tapejarid pterosaur

Huaxiadraco is a genus of tapejarid pterodactyloid pterosaur from the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China. It is the third valid genus of tapejarid from the Jehol Biota, after Sinopterus and Eopteranodon. It contains one species, Huaxiadraco corollatus, originally assigned to the defunct genus Huaxiapterus.

References

  1. ↑ David M. Martill; Mick Green; Roy Smith; Megan Jacobs; John Winch (2020). "First tapejarid pterosaur from the Wessex Formation (Wealden Group: Lower Cretaceous, Barremian) of the United Kingdom". Cretaceous Research. 113: Article 104487. doi:10.1016/j.cretres.2020.104487. S2CID   219099220.
  2. ↑ Pêgas, Rodrigo V. (2024-06-10). "A taxonomic note on the tapejarid pterosaurs from the Pterosaur Graveyard site (Caiuá Group, ?Early Cretaceous of Southern Brazil): evidence for the presence of two species". Historical Biology : 1–22. doi:10.1080/08912963.2024.2355664. ISSN   0891-2963.
  3. ↑ Peter Wellnhofer, Eric Buffetaut (1999). "Pterosaur remains from the Cretaceous of Morocco". Paläontologische Zeitschrift. 73 (1–2): 133–142. doi:10.1007/BF02987987. S2CID   129032233.
  4. ↑ Vullo, R.; Marugán-Lobón, J. S.; Kellner, A. W. A.; Buscalioni, A. D.; Gomez, B.; De La Fuente, M.; Moratalla, J. J. (2012). Claessens, Leon (ed.). "A New Crested Pterosaur from the Early Cretaceous of Spain: The First European Tapejarid (Pterodactyloidea: Azhdarchoidea)". PLOS ONE. 7 (7): e38900. Bibcode:2012PLoSO...738900V. doi: 10.1371/journal.pone.0038900 . PMC   3389002 . PMID   22802931.
  5. ↑ Mull, Olivia K.; Bennett, S. Christopher (2023-07-31). "Tapejarine pterosaur from the late Albian Paw Paw Formation of Texas, USA, with extensive feeding traces of multiple scavengers". Historical Biology. doi:10.1080/08912963.2023.2241044.
  6. ↑ Lü, J.; Jin, X.; Unwin, D.M.; Zhao, L.; Azuma, Y.; Ji, Q. (2006). "A new species of Huaxiapterus (Pterosauria: Pterodactyloidea) from the Lower Cretaceous of western Liaoning, China with comments on the systematics of tapejarid pterosaurs". Acta Geologica Sinica. 80 (3): 315–326. doi:10.1111/j.1755-6724.2006.tb00251.x. S2CID   129851866.
  7. 1 2 3 Pinheiro, F.L.; Fortier, D.C.; Schultz, C.L.; De Andrade, J.A.F.G.; Bantim, R.A.M. (2011). "New information on Tupandactylus imperator, with comments on the relationships of Tapejaridae (Pterosauria)". Acta Palaeontologica Polonica. 56: 567–580. doi: 10.4202/app.2010.0057 .
  8. 1 2 Eck, K.; Elgin, R.A.; Frey, E. (2011). "On the osteology of Tapejara wellnhoferi KELLNER 1989 and the first occurrence of a multiple specimen assemblage from the Santana Formation, Araripe Basin, NE-Brazil". Swiss Journal of Palaeontology . 130 (2): 277–296. doi:10.1007/s13358-011-0024-5. S2CID   84883165.
  9. ↑ Wu, Wen-Hao; Zhou, Chang-Fu; Andres, Brian (2017). "The toothless pterosaur Jidapterus edentus (Pterodactyloidea: Azhdarchoidea) from the Early Cretaceous Jehol Biota and its paleoecological implications". PLOS ONE. 12 (9): e0185486. Bibcode:2017PLoSO..1285486W. doi: 10.1371/journal.pone.0185486 . PMC   5614613 . PMID   28950013.
  10. ↑ Meng, X. (2008). "A New Species of Sinopterus from Jehol Biota and Reconstraction of Stratigraphic Sequence of the Jiufotang Formation". Thesis, Chinese Academy of Sciences.
  11. ↑ Could Tapejarid Pterosaurs be the dispersers of Klitzschophyllites angiosperm? A preliminary case of study of zoocory Flaviana J. Lima 1*, Renan A. M. Bantim1,2, Antônio A. F. Saraiva1 & Juliana M. Sayão3
  12. ↑ Mark P. Witton (2013), Pterosaurs: Natural History, Evolution, Anatomy, Princeton University Press, ISBN 978-0-691-15061-1
  13. ↑ Pêgas, R. V.; Costa, F. R.; Kellner, A. W. A. (2018). "New Information on the osteology and a taxonomic revision of the genus Thalassodromeus (Pterodactyloidea, Tapejaridae, Thalassodrominae)". Journal of Vertebrate Paleontology. 38 (2): e1443273. doi:10.1080/02724634.2018.1443273. S2CID   90477315.
  14. 1 2 3 Kellner, A.W.A.; Campos, D.A. (2007). "Short note on the ingroup relationships of the Tapejaridae (Pterosauria, Pterodactyloidea". Boletim do Museu Nacional. 75: 1–14.
  15. ↑ Lü, J., Unwin, D.M., Xu, L., and Zhang, X. (2008). "A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution." Naturwissenschaften,
  16. ↑ Martill, D.M., Bechly, G., and Heads, S.W. (2007). "Appendix: species list for the Crato Formation." In: Martill, D.M., Bechly, G., and Loveridge, R.F. (eds.), 2007. The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press, Cambridge. Pp. 582–607.
  17. 1 2 Martill, D.M.; Naish, D. (2006). "Cranial crest development in the azhdarchoid pterosaur Tupuxuara, with a review of the genus and tapejarid monophyly". Palaeontology. 49 (4): 925–941. doi: 10.1111/j.1475-4983.2006.00575.x .
  18. 1 2 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.
  19. ↑ Beccari, Victor; Pinheiro, Felipe Lima; Nunes, Ivan; Anelli, Luiz Eduardo; Mateus, Octávio; Costa, Fabiana Rodrigues (2021-08-25). "Osteology of an exceptionally well-preserved tapejarid skeleton from Brazil: Revealing the anatomy of a curious pterodactyloid clade". PLOS ONE. 16 (8): e0254789. doi: 10.1371/journal.pone.0254789 . ISSN   1932-6203. PMC   8386889 . PMID   34432814.
  20. ↑ 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 Reports9: 4940 doi : 10.1038/s41598-019-41280-4
  21. ↑ 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.
  22. ↑ Kellner, Alexander W. A.; Weinschütz, Luiz C.; Holgado, Borja; Bantim, Renan A. M.; Sayão, Juliana M. (19 August 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.
  23. ↑ Beccari, Victor; Pinheiro, Felipe Lima; Nunes, Ivan; Anelli, Luiz Eduardo; Mateus, Octávio; Costa, Fabiana Rodrigues (2021-08-25). "Osteology of an exceptionally well-preserved tapejarid skeleton from Brazil: Revealing the anatomy of a curious pterodactyloid clade". PLOS ONE. 16 (8): e0254789. doi: 10.1371/journal.pone.0254789 . ISSN   1932-6203. PMC   8386889 . PMID   34432814.
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