Tapejaridae

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Tapejarids
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") is a family of azhdarchoid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, [1] Spain, [2] the United States, [3] and China. The most primitive genera were found in China, indicating that the family has an Asian origin. [4]

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. [5] 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. [6] 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. [6]

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. [7] 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. [8] [9]

Classification

Life restorations of sinopterines Huaxiadraco and Sinopterus Jiufotang tapejarids.png
Life restorations of sinopterines Huaxiadraco and Sinopterus

Tapejaridae was named and defined by Brazilian paleontologist Alexander Kellner in 1989 as the clade containing both Tapejara and Tupuxuara, plus all descendants of their most recent common ancestor. In 2007, Kellner divided the family: Tapejarinae, consisting of Tapejara and its close relatives, and Thalassodrominae, consisting of Thalassodromeus and Tupuxuara. [10] A 2011 subsumed the family Chaoyangopterinae in as the subfamily Chaoyangopterinae, [5] something not followed by future authors. Kellner's concept of a Tapejaridae consisting of Tapejarinae and Thalassodrominae would be the basis for numerous subsequent phylogenetic analyses. [11] [12] [13] [14] [15]

Skeletal reconstructions of Thalassodromeus, phylogenetically controversial pterosaurs classically assigned to Tapejaridae Thalassodromeus in Japan.jpg
Skeletal reconstructions of Thalassodromeus , phylogenetically controversial pterosaurs classically assigned to Tapejaridae

Various opposing studies have arose challenging Kellner's concept of Tapejaridae. The 2003 model of paleontologist David Unwin found Tupuxara and Thalassodromeus to be more distantly related to Tapejara and therefore outside of Tapejaridae, instead being related to Azhdarchidae. [16] Later, in 2006, British paleontologists David Martill and Darren Naish followed Unwin's concept, and provided a revised definition for Tapejaridae was also proposed: the clade containing all species more closely related to Tapejara than to Quetzalcoatlus . [17] A 2008 study by Lü Junchang and colleagues also corroborated this model, and used the term "Tupuxuaridae" to include both genera. [18] In 2009, British paleontologist Mark Witton also agreed with the Unwin model. However, he noted that the term Thalassodrominae was created before Tupuxuaridae, meaning it had naming priority. He elevated Thalassodrominae to family level, thus creating the denomination Thalassodromidae. [19]

Regarding the core tapejarid clade, American paleontologist Brian Andres and colleagues formally defined Tapejaridae as the clade containing Tapejara and Sinopterus in 2014. They also re-defined the subfamily 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 . [20] In 2020, in the description of the genus Wightia , an opposing subfamily was named, Sinopterinae, consisting of tapejarids more closely related to Sinopterus than Tapejara. [21] These studies follow the Unwin model, opposing Kellner's model of Tapejaridae while corroborating the close relationship between thalassodromids, azhdarchids, rather than tapejarids.

Reconstructed skull of Caupedactylus, a pterosaur recovered either within Tapejarinae or just outside Tapejaridae in its own clade, Caupedactylia Unnamed pterosaur.jpg
Reconstructed skull of Caupedactylus , a pterosaur recovered either within Tapejarinae or just outside Tapejaridae in its own clade, Caupedactylia

In 2023, paleontologist Rodrigo Pêgas and colleagues argued that despite the disagreements about the position of Thalassodromeus and its relatives, the species in question were consistently related. Therefore, they favored the term Thalassodromidae to have consistency with other studies that used the same name, despite finding them to form a natural grouping with Tapejaridae in their phylogenetic analysis (per the Kellner model). Thus, Thalassodromidae and Tapejaridae would be separate families within Tapejaromorpha.In their 2023 study, Pêgas and colleagues redefined Tapejaridae to be the most recent common ancestor of Sinopterus, Tapejara, and Caupedactylus in order to preserve the scope of the family in light of finding Caupedactylus, traditionally a tapejarine, outside of the Andres definition of Tapejaridae. They divided this redefined Tapejaridae into the groups Eutapejaria, containing the subfamilies Sinopterinae and Tapejarinae, and Caupedactylia, containing the pterosaurs Caupedactylus and Aymberedactylus . [22] In 2024, Pêgas rejected this redefinition of Tapejaridae in light of non-compliance with phylocode rules, applying the Tapejara and Sinopterus definition and deeming Eutapejaria a synonym. Instead, he created the larger group contain Tapejaridae and Caupedactylia, removing Caupedactylus and Aymberedactylus from the family itself. [23]

The cladogram below shows the phylogenetic analysis conducted by paleontologist Gabriela Cerqueira and colleagues in 2021, which uses Kellner's nomenclature of Tapejaridae. [24]

Tapejaridae

Below are two cladograms representing different concepts of Tapejaridae. The first one shows the phylogenetic analysis conducted by Andres in 2021, in which Tapejaridae consists of the subfamilies Tapejarinae and Sinopterinae. He found the pterosaurs Lacusovagus and Keresdrakon as tapejarines, an arrangement that had never been recovered in previous analyses. Regarding the interrelationships of Tapejaridae, Andres follows Unwin's concept. [25] The second cladogram shows the phylogenetic analysis conducted by Pêgas in 2024. He also found Tapejaridae to consist of both Tapejarinae and Sinopterinae, but differed from Andres in recovering the tapejarid Bakonydraco as a sinopterine instead of tapejarine. He created the new subtribe Caiuajarina within Tapejarini to include Caiuajara and Torukjara . Additionally, his analysis further differs from that of Andres in finding both Tapejaridae and Thalassodromidae within Tapejaromorpha, which corroborates the close relationship between thalassodromids and tapejarids, similar to Kellner. [23]

Subclades

Summary of the phylogenetic definitons of tapejarid subclades as discussed in the classification section.

NameNamed byDefinitionNotes
CaiuajarinaPêgas, 2024 [26] The largest clade containing Caiuajara dobruskii , but not Tapejara wellnhoferi .
CaupedactyliaPêgas et al., 2023 [22] The largest clade containing Caupedactylus ybaka , but not Tapejara wellnhoferi .May be within Tapejaridae or just outside in the broader clade Tapejariformes.
EutapejariaPêgas et al., 2023 [22] The largest clade containing Tapejara wellnhoferi , but not Caupedactylus ybaka .May be synonymous with Tapejaridae when Caupedactylia is outside of Tapejaridae.
SinopterinaeLü et al., 2016The largest clade containing Sinopterus dongi , but not Tapejara wellnhoferi .
TapejaridaeKellner, 1989The smallest clade containing Tapejara wellnhoferi and Sinopterus dongi .Has had multiple interpretations of how inclusive the family is. Originally including Tapejara and relatives along with Thalassodromeus and relatives, then the last common ancestor and all descendants of Tapejara and Sinopterus, and most recently proposed as the last common ancestor Caupedactylus, Tapejara, and Sinopterus. The second interpretation is the currently most followed convention. [26]
TapejariformesPêgas et al., 2024 [26] The clade characterized by a downturned rostrum synapomorphic with that of Tapejara May be synonymous with Tapejaridae if Caupedactylus is a tapejarid.
TapejarinaeKellner & Campos, 2007 [10] The largest clade containing Tapejara wellnhoferi , but not Sinopterus dongi .Has been historically treated as one of two subfamilies (the other being Thalassodrominae) in Tapejaridae. More recently, it is treated as one of the two main subfamilies along with Sinopterinae.
TapejariniAndres et al., 2014 [20] The smallest clade containing Tapejara wellnhoferi and Tupandactylus imperator

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

Tupuxuara is a genus of thalassodromid pterosaur that lived during the Albian age of the Early Cretaceous, about 112 million years ago. Its remains were found in what is now the Romualdo Formation of the Santana Group in Brazil. Tupuxuara was named in 1988 by paleontologists Alexander Kellner and Diógenes de Almeida Campos. The name Tupuxuara means "familiar spirit" referring to a familiar spirit in the mythology of the Tupi people in Brazil. Two species have been named, T. longicristatus, the type species, and T. leonardii. An additional species has been named in 2013, T. deliradamus. However, the validity of this species has been put into question and it may not even belong to Tupuxuara.

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

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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

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<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

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  2. ↑ 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.
  3. ↑ 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.
  4. ↑ 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.
  5. 1 2 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 .
  6. 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.
  7. ↑ 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.
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  9. ↑ 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
  10. 1 2 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.
  11. ↑ 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.
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  15. ↑ 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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  17. ↑ 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. ↑ Lü J.; D.M. Unwin; Xu L.; Zhang X. (2008). "A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution". Naturwissenschaften. 95 (9): 891–7. doi:10.1007/s00114-008-0397-5. PMID   18509616.
  19. ↑ Witton, M. P. (2009). "A new species of Tupuxuara (Thalassodromidae, Azhdarchoidea) from the Lower Cretaceous Santana Formation of Brazil, with a note on the nomenclature of Thalassodromidae". Cretaceous Research. 30 (5): 1293–1300. Bibcode:2009CrRes..30.1293W. doi:10.1016/j.cretres.2009.07.006. S2CID   140174098.
  20. 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.
  21. ↑ Martill, David M.; Green, Mick; Smith, Roy; Jacobs, Megan; Winch, John (April 2020). "First tapejarid pterosaur from the Wessex Formation (Wealden Group: Lower Cretaceous, Barremian) of the United Kingdom" (PDF). Cretaceous Research. 113: 104487. doi:10.1016/j.cretres.2020.104487.
  22. 1 2 3 Pêgas, R. V.; Zhoi, X.; Jin, X.; Wang, K.; Ma, W. (2023). "A taxonomic revision of the Sinopterus complex (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota, with the new genus Huaxiadraco". PeerJ. 11. e14829. doi: 10.7717/peerj.14829 . PMC   9922500 .
  23. 1 2 Pêgas, Rodrigo V. (June 10, 2024). "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.
  24. ↑ Cerqueira GM, Santos MA, Marks MF, Sayão JM, Pinheiro FL (2021). "A new azhdarchoid pterosaur from the Lower Cretaceous of Brazil and the paleobiogeography of the Tapejaridae". Acta Palaeontologica Polonica. 66. doi: 10.4202/app.00848.2020 ..
  25. ↑ Andres, Brian (December 7, 2021). "Phylogenetic systematics of Quetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea)". Journal of Vertebrate Paleontology. 41 (sup1): 203–217. Bibcode:2021JVPal..41S.203A. doi: 10.1080/02724634.2020.1801703 . ISSN   0272-4634. S2CID   245078533.
  26. 1 2 3 Pêgas, Rodrigo V. (2024). "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.
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