''Santanadactylus'' (meaning "

Santana Formation 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 sta ...

finger") was a

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 ...

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 to ...

from the

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 ...

-age Romualdo Member of the

Upper Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...

Santana Formation, of Barra do Jardim,

Araripe Plateau The Chapada do Araripe, also known as the Serra do Araripe, is a ''chapada'' (plateau) in northeastern Brazil. The chapada forms the boundary of Ceará and Pernambuco states, and forms the watershed between the Jaguaribe River of Ceará, which flow ...

, Ceará State,

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 ...

. Four species have been named, but today are no considered congeneric with each other. It was a rather large pterosaur.

History and species

The genus was named in 1980 by the Dutch

paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...

Paul de Buisonjé. 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 ''S. brasilensis'', the specific name referring to Brazil. It is based on

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 ...

UvA M 4894 ( Geological Institute of the University of Amsterdam), an upper part of the right

humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...

and a right

scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...

coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...

. UvA M 4895, consisting of two

cervical In anatomy, cervical is an adjective that has two meanings: # of or pertaining to any neck. # of or pertaining to the female cervix: i.e., the ''neck'' of the uterus. *Commonly used medical phrases involving the neck are **cervical collar **cerv ...

vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...

e from a different individual, was assigned as a paratype, referred because it was found in the same lot of 25 chalk nodules bought from collectors.De Buisonjé, P.H. (1980). ''Santanadactylus brasilensis'' nov. gen. nov. sp. a longnecked, large pterosaur from the Aptian of Brazil. ''Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen'' B 83(2):145-172. Additional remains, including a

notarium Notarium or os dorsale is a bone consisting of the fused vertebra of the shoulder in birds and some pterosaurs. The structure helps brace the chest against the forces generated by the wings. In birds, the vertebrae are only in contact with adjac ...

(fused vertebrae supporting the

shoulder The human shoulder is made up of three bones: the clavicle (collarbone), the scapula (shoulder blade), and the humerus (upper arm bone) as well as associated muscles, ligaments and tendons. The articulations between the bones of the shoulder mak ...

) were subsequently referred to the taxon by Wellnhofer ''et al.'' (1983) and Wellnhofer (1991).Wellnhofer, P., Buffetaut, E., and Gigase, P. (1983). A pterosaurian notarium from the Lower Cretaceous of Brazil. ''Paläontologische Zeitschrift'' 57147-157.Wellnhofer, P. (1991). Weitere Pterosaurierfunde aus der Santana-Formation (Apt) der Chapada do Araripe, Brasilien. ''Palaeontographica'' A 215:43–101.

erman Erman Rašiti may refer to: Given name * Erman Bulucu (born 1989), Turkish footballer * Erman Eltemur (born 1993), Turkish karateka * Erman Güraçar (born 1974), Turkish footballer * Erman Kılıç (born 1983), Turkish footballer * Erman Kunter (b ...

/ref> In 1985,

Peter Wellnhofer Peter Wellnhofer (born Munich, 1936) is a German paleontologist at the Bayerische Staatssammlung fur Paläontologie in Munich. He is best known for his work on the various fossil specimens of ''Archaeopteryx'' or "Urvogel", the first known bird. W ...

, a German paleontologist who has written numerous

scientific publication : ''For a broader class of literature, see Academic publishing.'' Scientific literature comprises scholarly publications that report original empirical and theoretical work in the natural and social sciences. Within an academic field, scient ...

s on pterosaurs, named three additional species: ''S. araripensis'', ''S. pricei'', and ''S. spixi''.Wellnhofer, P. (1985). Neue Pterosaurier aus der Santana-Formation (Apt) der Chapada do Araripe, Brasilien. ''Paläontographica'' A 187:105-182.

/ref> ''S. araripensis'', named after the

, was a large species based on BSP 1982 I 89, remains including a partial skull (missing the end of the jaws) and arms; the preserved skull section had no crest. ''S. pricei'', named after

Llewellyn Ivor Price Llewellyn Ivor Price (October 9, 1905 – June 9, 1980) was one of the first Brazilian paleontologists. His work contributed not only to the development of Brazilian but also to global paleontology. He collected '' Staurikosaurus'', the first ...

, was the smallest of the three species; it was based on BSP 1980 I 122, a left wing from the

elbow The elbow is the region between the arm and the forearm that surrounds the elbow joint. The elbow includes prominent landmarks such as the olecranon, the cubital fossa (also called the chelidon, or the elbow pit), and the lateral and the media ...

down, and additional arm material has been referred to it over the years. ''S. spixi'', intermediate in size, was based on BSP 1980 I 121, another left wing, the name honoring

Johann Baptist von Spix Johann Baptist Ritter von Spix (9 February 1781 – 13 March 1826) was a German natural history, biologist. From his expedition to Brazil, he brought to Germany a large variety of specimens of plants, insects, mammals, birds, amphibians and fish. ...

. Over the years, the species of this taxon have been reassessed. Chris Bennett suggested that the hypodigm of ''S. brasilensis'' was a

chimera Chimera, Chimaera, or Chimaira (Greek for " she-goat") originally referred to: * Chimera (mythology), a fire-breathing monster of Ancient Lycia said to combine parts from multiple animals * Mount Chimaera, a fire-spewing region of Lycia or Cilicia ...

of a pteranodontid and something else (in that the holotype and paratype belonged to different forms), ''S. araripensis'' and ''S. pricei'' were pteranodontids, and ''S. spixi'' was a

dsungaripterid Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea. They were robust pterosaurs with good terrestrial abilities and flight honed for inland settings. Classification In 1964 Young created a family to place the recentl ...

.Bennett, S.C. (1989). A pteranodontid pterosaur from the Early Cretaceous of Peru, with comments on the relationships of Cretaceous pterosaurs. ''Journal of Paleontology'' 63:669-677. Wellnhofer (1991) removed ''S. spixi'' from the genus as well. Kellner and Campos (1992) agreed that ''S. spixi'' was not congeneric with ''S. brasiliensis'', but suggested that it was a

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 primit ...

.Kellner, A.W.A., and Campos, D. de A. (1992). A new tapejarid from the Santana Formation (Lower Cretaceous) from the Araripe Basin, Northeastern Brasil. ''Journal of Vertebrate Paleontology'' 12(Supplement to 3):36–37A. Unwin (2003) proposed that "Santanadactylus" ''spixi'' was a species of ''

Tupuxuara ''Tupuxuara'' is a genus of large, crested, and toothless Pterodactyloidea, pterodactyloid pterosaur from the Early Cretaceous period (Albian stage) of what is now the Romualdo Formation of the Santana Group, Brazil, about 125 to 112 million ye ...

'' based on comparison with specimens of this genus. Averianov (2014) considered ''Santanadactylus spixi'' a ''nomen dubium'' probably synonymous with ''Tupuxuara longicristatus'', and he also indicated that the paratype of ''S. brasiliensis'' was likely referable to ''T. longicristatus''. Kellner (1990) renamed ''S. araripensis'' to ''

Anhanguera Anhanguera may refer to: People * Bartolomeu Bueno da Silva (1672–1740), a bandeirante Places in Brazil * Anhanguera, Goiás, a municipality in the state of Goiás * Anhanguera (district of São Paulo), a district in São Paulo * Parque Anhan ...

araripensis'', followed by Wang ''et al.'' (2008), though Veldmeijer (2003) included it in ''

Coloborhynchus ''Coloborhynchus'' is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. ''Coloborhynchus'' is known from the Lower Cretaceous of Engl ...

''. Recent study, however, considers ''S. araripensis'' dubious.

Classification

De Buisonjé first assigned ''Santanadactylus'' to Criorhynchidae (=Ornithocheiridae). Wellnhofer (1991) considered it a member of the

Ornithocheiridae Ornithocheiridae (or ornithocheirids, meaning "bird hands") is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the Early to Late Cr ...

based on the structure of the humerus, but noted that the elongation of the paratype neck vertebrae distinguished it from other ornithocheirids. Bennett (1989) assigned ''S. brasilensis'' (holotype only) to

Pteranodontidae The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the rear of the head ( ...

. Kellner (1990), concluding that he could find but a single

autapomorphy 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 ...

for ''Santanadactylus brasilensis'', the straight ventral margin of the proximal part of the deltopectoral crest, assigned the genus to Pterodactyloidea ''

incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...

'', but later (Kellner and Tomida 2000) assigned it to

Pteranodontoidea Pteranodontoidea (or pteranodontoids, from Greek meaning "toothless wings") is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous (early Valanginian to late Maastrichtian stages) of Asia, Africa, Europe, North Americ ...

. ''S. pricei'' according to Kellner belonged to a

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

descending from the last common ancestor of ''

Istiodactylus ''Istiodactylus'' is a genus of pterosaur that lived during the Early Cretaceous period, about 120 million years ago. The first fossil was discovered on the English Isle of Wight in 1887, and in 1901 became the holotype specimen of a new species ...

'' and the

Anhangueridae Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a later ...

. The same was in his analysis true for ''

Araripesaurus ''Araripesaurus'' is a genus a pterosaur belonging to the suborder Pterodactyloidea, it was discovered in the Romualdo Formation of the Santana Group in northeastern Brazil, which dates back to the Aptian and Albian of the Early Cretaceous. The ...

'', a genus of which he had previously thought ''S. pricei'' was a

junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...

Paleobiology

''Santanadactylus'' is regarded as a large pterosaur, Wellnhofer for the various species indicating a

wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan of ...

of 2.9–5.7 meters (9.5–18.7 ft). De Buisonjé thought ''Santanadactylus brasilensis'' had a wingspan of .Buisonjé, P. H. de. (1981) "''Santanadactylus brasilensis'': Skelet-reconstructie van een vliegend reptiel met zes meter vlucht", ''Gea'', 14(2): 37-48 It may have been adapted for gliding over flapping

flight Flight or flying is the process by which an object moves through a space without contacting any planetary surface, either within an atmosphere (i.e. air flight or aviation) or through the vacuum of outer space (i.e. spaceflight). This can be a ...

.Glut, D.F. (2006). ''Santanadactylus'' In: Glut, D.F. ''Dinosaurs: The Encyclopedia.'' Supplement 4. McFarland & Company, Inc.:Jefferson, North Carolina, 628-629. .

References

External links

''Santanadactylus''
in The Pterosauria. Accessed 2007–02–09

a posting from George Olshevsky on the Dinosaur Mailing List, which, although incomplete, should give some idea as to the complexity of the taxonomy here. Accessed 2007–02–09 {{Portal bar, Paleontology, Cretaceous, Brazil Pteranodontoids Early Cretaceous pterosaurs of South America Cretaceous Brazil Fossils of Brazil Albian life Fossil taxa described in 1980

History and species

Classification

Paleobiology

See also

References

External links