''Tapejara'' (from a Tupi word meaning "the lord of the path" https://www.teses.usp.br/teses/disponiveis/41/41133/tde-14082021-135310/publico/Lucas_Piazentin_vol_II_CORRIG_compac.pdf) is a

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...

ian

pterosaur Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...

from the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

Period ( Santana Group, dating to about 127 to 112 million years ago). ''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 .

History of discovery

Araripe_Basin_map_-_formations_and_resources

The

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

of ''Tapejara'' (MN 6595-V, originally catalogued as CD-R 080) consists of a partial skull. It was described in 1989 by Alexander W. A. Kellner, who recognised it as a new genus and species. Kellner named it ''Tapejara wellnhoferi'', ostensibly translating to "Wellnhofer's old being". However, the name actually derives from the Tupí-Guaraní ''tape'' ("path") and ''jara'' ("lord"), and therefore, ''T. wellnhoferi'' actually translates to "Wellnhofer's lord of the path". Another, more complete specimen, AMNH 24440, was discovered in the same year as Kellner's original description, in a

calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime (mineral), lime or being chalky. The term is used in a wide variety of Science, scientific disciplines. In zoology ''Calcare ...

concretion A concretion is a hard and compact mass formed by the precipitation of mineral cement within the spaces between particles, and is found in sedimentary rock or soil. Concretions are often ovoid or spherical in shape, although irregular shapes a ...

. The providence of this specimen is unclear, though it likely comes from either the Jardim or Santana do Cariri localities. AMNH 24440 consists of a partial skull, a partial

mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...

(lower jaw), and an

anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...

(front) cervical (neck)

vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...

. In 2011, another, SMNK PAL 1137, was described, also found in a concretion. This specimen consists of a fairly complete premaxillo-maxilla, a unit consisting of the fused premaxillae and maxillae, and much of the

braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calv ...

. The locality from which it originates is uncertain, though given that the concretion was originally blue, it likely comes from the Sierra de Maosina locality, part of the Romualdo Formation of the Santana Group.

Species and classification

The type species and only one currently recognized as valid by most researchers, is ''T. wellnhoferi''. The specific name honors German paleontologist Peter Wellnhofer. Two larger species, originally named ''Tapejara imperator'' and ''Tapejara navigans'', were later also placed in the genus ''Tapejara'' upon discovery. However, several studies have shown that ''T. imperator'' and ''T. navigans'' are significantly different from ''T. wellnhoferi'' and therefore were reclassified into new genera. The species ''T. imperator'' was given its own genus, '' Tupandactylus'', 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 Diogenes de Almeida Campos. Unwin and Martill found that ''T. imperator'' and ''T. navigans'' belong in the same genus, and named them ''Ingridia imperator'' and ''I. navigans'', respectively. This genus name honored Wellnhofer's late wife Ingrid.Unwin, D. M. and Martill, D. M. (2007). "Pterosaurs of the Crato Formation." In Martill, D. M., Bechly, G. and Loveridge, R. F. (eds), ''The Crato Fossil Beds of Brazil: Window into an Ancient World.'' Cambridge University Press (Cambridge), pp. 475–524. Because ''Tupandactylus'' was named first, it retained priority over the name ''Ingridia''. To complicate matters, both the name ''Tupandactylus'' and ''Ingridia'' used the former ''Tapejara imperator'' as their

type species In International_Code_of_Zoological_Nomenclature, 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 spe ...

.Naish, D. (2008). "Crato Formation fossils and the new tapejarids." Weblog entry. ''Tetrapod Zoology''. January 18, 2008. Accessed January 31, 2008 (). The scientists who described ''Tupandactylus'' did not name a ''Tupandactylus navigans'' (but instead suggested it was synonymous to ''Tupandactylus imperator''), and ''Tapejara navigans'' was not formally reclassified as a distinct species of ''Tupandactylus'' until 2011. Cast of Tapejara wellnhoferi - Pterosaurs Flight in the Age of Dinosaurs

Cast of Tapejara wellnhoferi - Pterosaurs Flight in the Age of Dinosaurs

The cladogram below follows a

phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...

by Kellner, the describer of ''Tapejara'', and colleagues in 2019. They recovered ''Tapejara'' within the Tapejarini (a tribe within the family Tapejaridae), sister taxon to three other genera: '' Europejara'', '' Caiuajara'', and '' Tupandactylus''.

Paleobiology

Comparisons between the scleral rings of ''Tapejara'' and modern birds and reptiles suggest that it may have been cathemeral, active throughout the day at short intervals.

References

{{Portal bar, Paleontology, Brazil Tapejaromorphs Early Cretaceous pterosaurs of South America Albian life Aptian life Cretaceous Brazil Fossils of Brazil Romualdo Formation Fossil taxa described in 1989 Taxa named by Alexander Kellner Tupi–Guarani languages

History of discovery

Species and classification

Paleobiology

See also

References