24-640: Torukjara bandeirae , Caiuajara dobruskii , Tupandactylus imperator , Tapejara wellnhoferi , Huaxiadraco corollatus , and Sinopterus dongi 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 , Spain , the United States , and China . The most primitive genera were found in China, indicating that
48-424: 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 . Andres and colleagues did not follow this proposal, instead formally defining Tapejaridae as
72-546: A new genus and species of tapejarid pterosaurs based on these fossil remains. The generic name , Torukjara , combines a reference to the fictional toruk creature of the Avatar media franchise with the Tupi word " jara ", meaning "lord", a suffix often used for tapejarid genera. The specific name , bandeirae , honours Kamila Bandeira, a Brazilian palaeontologist who first studied the possibility that multiple species could be present in
96-1099: 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. Azhdarchids are characterized by their long legs and extremely long necks, made up of elongated neck vertebrae which are round in cross section. Most species of azhdarchids are still known mainly from their distinctive neck bones and not much else. The few azhdarchids that are known from reasonably good skeletons include Zhejiangopterus and Quetzalcoatlus . Azhdarchids are also distinguished by their relatively large heads and long, spear-like jaws. There are two major types of azhdarchid morphologies:
120-649: Is a stub . You can help Misplaced Pages by expanding it . Azhdarchidae Azhdarchidae (from the Persian word azhdar , اژدر , a dragon-like creature in Persian mythology ) 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 (late Berriasian age, about 140 million years ago). Azhdarchids are mainly known for including some of
144-432: Is an extinct genus of tapejarin pterodactyloid pterosaurs from the Early Cretaceous "Pterosaur Graveyard" ( Caiuá Group ) of Brazil. The genus contains a single species , T. bandeirae , known from several specimens. The Torukjara fossil material was originally assigned to Caiuajara , but several anatomical differences indicate that both similar species coexisted in the ecosystem. The Torukjara fossil material
168-764: Is currently unnamed, recently discovered in Transylvania , may be the largest representative of the family thus far discovered. This unnamed specimen (nicknamed " Dracula " by paleontologists), currently on display in the Altmühltal Dinosaur Museum in Bavaria is estimated to have a wingspan of 12–20 m (39–66 ft), although similarities to the contemporary azhdarchid Hatzegopteryx have also been noted. Azhdarchids were originally classified as close relatives of Pteranodon due to their long, toothless beaks. Others have suggested they were more closely related to
192-481: 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 . Assuming thalassodromines are tapejarids, this clade also included raptorial, macropredatory species adapted to hunt proportionally large prey. Tapejaridae
216-1073: Is the one by Kellner and colleagues in 2019. The cladogram on the right shows their phylogenetic analysis . In 2021 a new study focus on the most complete Tapejaridae, a skeleton Tupandactylus navigans , confirmed the 2019 topology . Topology 1 : Andres and colleagues (2014). Neoazhdarchia Bennettazhia Eopteranodon lii "Sinopterus" gui Nemicolopterus crypticus Huaxiapterus jii Sinopterus dongi "Huaxiapterus" benxiensis "Huaxiapterus" corollatus Tupandactylus navigans Tupandactylus imperator Bakonydraco galaczi Europejara olcadesorum Tapejara wellnhoferi Topology 2 : Kellner and colleagues (2019). Keresdrakon Thalassodromeus Tupuxuara Caupedactylus Aymberedactylus Eopteranodon "Huaxiapterus" benxiensis "Huaxiapterus" corollatus Sinopterus Europejara Caiuajara Tapejara Tupandactylus [REDACTED] [REDACTED] [REDACTED] Torukjara bandeirae Torukjara
240-549: The "blunt-beaked" forms with shorter and deeper bills and the "slender-beaked" forms with longer and thinner jaws. It had been suggested azhdarchids were skimmers, but further research has cast doubt on this idea, demonstrating that azhdarchids lacked the necessary adaptations for a skim-feeding lifestyle, and that they may have led a more terrestrial existence similar to modern storks and ground hornbills . Most large azhdarchids probably fed on small prey, including hatchling and small dinosaurs; in an unusual modification of
264-477: 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 (in the clade Neoazhdarchia ). 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
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#1732851340529288-506: The azhdarchid bodyplan, the robust Hatzegopteryx may have tackled larger prey as the apex predator in its ecosystem. In another departure from typical azhdarchid lifestyles, the jaw of Alanqa may possibly be an adaptation to crushing shellfish and other hard foodstuffs. Azhdarchids are generally medium- to large-sized pterosaurs, with the largest achieving wingspans of 10–12 metres (33–39 ft), but several small-sized species have recently been discovered. Another azhdarchid that
312-455: 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 . Below are two alternate cladograms: the first, presented by Andres and colleagues in 2014, found the a grouping of tapejarids at the base of
336-459: 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. 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. An example of one of these studies
360-414: The family has an Asian origin. 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
384-581: The fossil material. In their phylogenetic analysis , Pêgas (2024) recovered Torukjara as a tapejarine pterosaur in the family Tapejaridae, as the sister taxon to Caiuajara . The clade containing Torukjara and Caiurajara was named as the new subtribe Caiuajarina. These results are displayed in the cladogram below: Thalassodromidae Caupedactylia Sinopterinae Europejara Tupandactylus spp. Tapejara Caiuajara Torukjara [REDACTED] [REDACTED] [REDACTED] This pterosaur -related article
408-462: 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 (now known as Arambourgiania ). They were among the last known surviving members of the pterosaurs, and were a rather successful group with
432-568: 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. 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. Tapejarids had unusually reduced shoulder girdles that would have been slung low on
456-439: The tapejarines than to the thalassodromines. Felipe Pinheiro and colleagues (2011) reclassified the group as a subfamily of Tapejaridae, Chaoyangopterinae , for this reason. 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
480-467: The thalassodromines are more closely related to the azhdarchids 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 , often found to be closer to azhdarchids, represent a lineage within the Tapejaridae, more closely related to
504-640: The toothy ctenochasmatids (which include filter-feeders like Ctenochasma and Pterodaustro ). Currently it is widely agreed that azhdarchids were closely related to pterosaurs such as Tupuxuara and Tapejara . Classification after Unwin 2006, except where noted. One of the most complete cladograms of azhdarchids is presented by Andres (2021): Azhdarcho Albadraco Aerotitan Mistralazhdarcho Phosphatodraco Aralazhdarcho Eurazhdarcho Zhejiangopterus Wellnhopterus Cryodrakon Hatzegopteryx Arambourgiania Quetzalcoatlus [REDACTED] In
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#1732851340529528-503: The torso, resulting in wings that protruded from near the belly rather than near the back, a "bottom decker" arrangement reminiscent of some planes. 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. Direct evidence for plant-eating
552-554: 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 . Some studies, such as one by Lü and colleagues in 2008, have found that
576-562: Was discovered in the Pterosaur Graveyard locality which likely belongs to the Rio Paraná Formation ( Caiuá Group ) of Cruzeiro do Oeste , Paraná State , Brazil. The holotype specimen (CP.V 8175), discovered in 2014, consists of a partial skeleton including a well-preserved skull. This specimen was first described in 2022 as belonging to the coeval Caiuajara . In 2024, Rodrigo V. Pêgas described Torukjara bandeirae as
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