''Caupedactylus'' is an

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

tapejarid 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 primiti ...

azhdarchoid 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 ...

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 to ...

known from the

Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma. Geology Pro ...

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 ± 0.9 M ...

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

Araripe Basin The Araripe Basin () is a rift basin covering about ,Neto et al., 2013, p.1 in Ceará, Piauí and Pernambuco states of northeastern Brazil. It is bounded by the Patos and Pernambuco lineaments, and is situated east of the Parnaíba Basin, southwes ...

in northeastern

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 ...

. The type and only species of ''Caupedactylus'' is ''Caupedactylus ybaka''.

Discovery and naming

The genus was first named and described by Alexander Wilhelm Armin Kellner in

2013 File:2013 Events Collage V2.png, From left, clockwise: Edward Snowden becomes internationally famous for leaking classified NSA wiretapping information; Typhoon Haiyan kills over 6,000 in the Philippines and Southeast Asia; The Dhaka garment fact ...

. 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 specimen ...

is ''Caupedactylus ybaka''. The generic name combines the goddess of beauty of the

Tupi Tupi may refer to: * Tupi people of Brazil * Tupi or Tupian languages, spoken in South America ** Tupi language, an extinct Tupian language spoken by the Tupi people * Tupi oil field off the coast of Brazil * Tupi Paulista, a Brazilian municipalit ...

, Caupe, with a Greek δάκτυλος, ''daktylos'', "finger", a usual suffix in the names of pterosaurs. The

specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...

''ybaka'' means "dwelling in the sky" in the local Tupinambá. ''Caupedactylus'' is known from 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 several ...

specimen MN 4726-V, probably found in the Araripe basin in a layer of the Romualdo Formation dating from the

, about 110 million years old. It consists of a partial skeleton with a complete

skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...

lower jaw In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...

s and a partial

postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...

l skeleton. The postcrania contain the right shoulder girdle, left coracoid, sternum, right humerus and the proximal part of the first phalanx of the wing finger. It represents and adult individual, which is exceptional for pterosaurs from this formation. The specimen was freed from the chalkstone by an acid bath.

Description

Kellner estimated the wingspan of ''Caupedactylus'' at , making it the largest known tapejarine. The skull is about forty-six centimetres long. Some unique traits,

autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...

, were established. The snout sports a crest on the

premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...

extending both to the front and to the rear of the skull. The point of the snout curves downwards combined with a slightly concave lower edge of the upper jaw. The paired points of the snout are not separated. The ascending branch of the jugal bone, connecting to the lacrimal bone, is strongly reclining at an angle of 115 degrees. In the rear of the palate a reduced split-shaped ''fenestra postpalatina'' is present. The quadratum reclines 150 degrees relative to the lower edge of the skull. The basisphenoid of the lower braincase lacks appending processes. The basisphenoid does not reach the space between the pteroids. The rear half of the lower edge of the symphysis of the lower jaws is thick, forming a slightly convex surface. The skull is crowned by a large rounded crest that continues to behind, descending over the parietals. The crest is very thin, about half a millimetre at the upper part. The jaws are toothless. The profile of the symphysis of the lower jaws closely fits the edge of the upper jaws: when closing the beak, no gap would have remained. The symphysis has a low keel at the underside. The palate is well-preserved and seems to show that the traditional interpretation of its bones is the correct one, contra the more modern view that what have usually been seen as the fronts of the palatine bones would really be the internal wings of the

maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...

Classification

''Caupedactylus'' was, 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 primiti ...

, placed in the subfamily

Tapejarinae 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 primit ...

, though without a

cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...

analysis. In 2019 however, Kellner (the describer of ''Caupedactylus'') and colleagues had included it in a

phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

where it was recovered as the

basalmost In phylogenetics, basal is the direction of the ''base'' (or root) of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to th ...

tapejarine.

References

{{Portal bar, Paleontology, Cretaceous, Brazil Tapejaromorphs Early Cretaceous pterosaurs of South America Cretaceous Brazil Fossils of Brazil Fossil taxa described in 2013 Taxa named by Alexander Kellner

Discovery and naming

Description

Classification

See also

References