''Tupuxuara'' is a

genus 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 nom ...

of large, crested, and toothless

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 ...

pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...

from the Early Cretaceous period (

Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/ Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0 ...

stage) of what is now the

Romualdo Formation The Romualdo Formation is a geologic Konservat-Lagerstätte in northeastern Brazil's Araripe Basin where the states of Pernambuco, Piauí and Ceará come together. The geological formation, previously designated as the Romualdo Member of the Sa ...

of the

Santana Group The Santana Group is a geologic group, formerly included as the middle part of the Araripe Group, in the Araripe Basin of northeastern Brazil. The group comprises the Crato, Ipubi and Romualdo Formations and is dated to the Aptian to Albian stag ...

Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...

, about 125 to 112 million years ago. ''Tupuxuara'' is a close relative of ''

Thalassodromeus ''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 collect ...

'', and both form a group that is either called

Thalassodrominae Thalassodrominae or Thalassodromidae (meaning "sea runners", due to previous misconceptions of skimming behavior; they are now thought to be terrestrial predators) is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional ...

(if placed within the family

Tapejaridae Tapejaridae (from a Tupi word meaning "the old being") 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 primi ...

) or Thalassodromidae (if placed within the clade

Neoazhdarchia Azhdarchoidea (or azhdarchoids) 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 o ...

Discovery

Tupuxuara leonardii Exhibit Museum of Natural History

The genus was named and described by

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 ...

and Diógenes de Almeida Campos in 1988.Kellner, A.W.A., and Campos, D.A. (1988). "Sobre un novo pterossauro com crista sagital da Bacia do Araripe, Cretaceo Inferior do Nordeste do Brasil. (Pterosauria, Tupuxuara, Cretaceo, Brasil)." ''Anais da Academia Brasileira de Ciências'', 60: 459–469.

n Portuguese N, or n, is the fourteenth Letter (alphabet), letter in the Latin alphabet, used in the English alphabet, modern English alphabet, the alphabets of other western European languages and others worldwide. Its name in English is English alphabet# ...

/ref> The

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...

is ''Tupuxuara longicristatus''. The generic name refers to a

familiar spirit In European folklore of the medieval and early modern periods, familiars (sometimes referred to as familiar spirits) were believed to be supernatural entities that would assist witches and cunning folk in their practice of magic. According to ...

from the

mythology Myth is a folklore genre consisting of narratives that play a fundamental role in a society, such as foundational tales or origin myths. Since "myth" is widely used to imply that a story is not objectively true, the identification of a narra ...

of the Tupi. The specific name means "long-crested" in

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

. The

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...

, MN 6591-V, was found in the Early Cretaceous (

) Santana Formation (now known as the

) of Brazil. It consists of a snout and some partial wing bones. Mature individuals of ''T. longicristatus'' had a back-swept crest arising from the snout. Much more fossil material has later been found, showing considerable variation in

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...

. Some researchers explain this as intra-specific variability, being caused by a difference in age or sex. Others, however, assume there are different species present. The largest specimens indicate a wingspan of , body length of and body msss of . In 1994 a second species was named by Kellner: ''Tupuxuara leonardii''. The specific name honors Giuseppe Leonardi. The holotype is MN 6592-V, a fragmentary skull with a more rounded crest. Other such material has been referred to ''T. leonardii''. Tupuxuara leonardii size

In 2009 a third species was named, by Mark Witton: ''Tupuxuara deliradamus''. The holotype is SMNK PAL 6410, a skull. Another skull is the

paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...

: KPMNH DL 84. The specific name is derived from Latin ''delirus'', "insane" or "crazy", and ''adamas'', "invincible", but also the word from which "diamond" is derived. The species has a distinctive diamond-shaped skull opening and low eye sockets. The name is a tribute to the song "

Shine On You Crazy Diamond "Shine On You Crazy Diamond" is a nine-part Pink Floyd composition written by David Gilmour, Roger Waters, and Richard Wright. It appeared on Pink Floyd's 1975 concept album ''Wish You Were Here''. The song is written about and dedicated to Syd B ...

" by Pink Floyd, one of Witton's favorite bands.

Classification

''Tupuxuara'' is a member of the group

Azhdarchoidea Azhdarchoidea (or azhdarchoids) 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 ...

. Kellner assigned it to the

within Azhdarchoidea. According to some analyses however, ''Tupuxuara'' is closer to the Azhdarchidae (the group that includes the giant Texan form '' Quetzalcoatlus'') than to '' Tapejara'' and its relatives. The cladogram below follows the 2011 analysis of Felipe Pinheiro and colleagues. Below is a

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 ...

showing the phylogenetic placement of ''Tupuxuara'' within Neoazhdarchia from Andres and Myers (2013).

Paleobiology

It was once been suggested that ''Tupuxuara'' was a fish eater at the coasts of

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...

, while some deviant hypotheses include the possibility it was a fruit eater. However, based on its azhdarchoid affinities, it was most likely a terrestrial omnivore or carnivore. The closely related ''

'' was specialized for larger prey, while both ''Tupuxuara'' species lacked such specializations. A subadult described by David Martill and

Darren Naish Darren William Naish is a British vertebrate palaeontologist, author and science communicator. As a researcher, he is best known for his work describing and reevaluating dinosaurs and other Mesozoic reptiles, including '' Eotyrannus'', '' Xenop ...

from the University of Portsmouth in 2006 had not yet fully developed its crest, which supports the suggestion that the crest was a marker for sexual maturity. Comparisons between the scleral rings of ''Tupuxuara'' and modern birds and reptiles suggest that it may have been diurnal.

References

External links

BBC News, "Flying reptile mystery 'solved'"
July 27, 2006 {{Portal bar, Paleontology, Cretaceous, Brazil Tapejaromorphs Early Cretaceous pterosaurs of South America Cretaceous Brazil Fossils of Brazil Crato Formation Romualdo Formation Fossil taxa described in 1988 Taxa named by Alexander Kellner Extinct reptiles

Discovery

Classification

Paleobiology

See also

References

External links