Santanadactylus
Encyclopedia
Santanadactylus was a genus
of pterodactyloid pterosaur
from the Aptian
-age Romualdo Member of the Lower Cretaceous Santana Formation, of Barra do Jardim, Araripe Plateau, Ceará Province
, Brazil
. Four species have been named, but today it is doubted they are part of the same genus. It was a rather large pterosaur.
is S. brasilensis, the specific name referring to Brazil. It is based on holotype
UvA M 4894 (Geological Institute of the University of Amsterdam), an upper part of the right humerus
and a right scapula
coracoid; other remains were also included in the genus: this paratype
consisted of two cervical
vertebrae from a different individual, referred because they were found in the same lot of 25 chalk nodules bought from collectors. Additional remains, including a notarium
(fused vertebrae supporting the shoulder
) have been referred to it since then. The structure of the humerus suggests an ornithocheirid
, but the long neck vertebrae argue against this.
In 1985, Peter Wellnhofer
, a German paleontologist who has written numerous scientific publications on pterosaurs, named three additional species: S. araripensis, S. pricei, and S. spixi. S. araripensis, named after the Araripe Plateau, 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
, was the smallest of the three species; it was based on BSP 1980 I 122, a left wing from the elbow
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 honouring Johann Baptist von Spix
.
Over the years, the species of this taxon have been reassessed. Chris Bennett suggested that S. brasilensis was a chimera
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
. Wellnhofer removed S. spixi from the genus as well, in 1991. In 1992, Alexander Kellner and Diogenes de Almeida Campos suggested that S. spixi was a tapejarid
; David Unwin in 2003 thought that "Santanadactylus" spixi was a species of Tupuxuara
. Kellner in 1990 renamed S. araripensis to Anhanguera
araripensis, followed by Wang and colleagues in 2008, though Veldmeijer (2003) included it in Coloborhynchus
. Part of the problem is the complicated taxonomy
of Santana Formation pterosaurs and their English
contemporaries, involving numerous genera, such as Amblydectes, Anhanguera, Araripesaurus
, Criorhynchus, Coloborhynchus, Lonchodectes
, Ornithocheirus
, and Tropeognathus.
Also the phylogeny is contentious. De Buisonjé first placed the genus into the Criorhynchidae, a concept that is no longer used. Wellnhofer considered it a member of the Ornithocheiridae
. According to Bennett S. brasilensis belonged to Pteranodontidae
sensu Bennett. Kellner, concluding that he could find but a single autapomorphy
for Santanadactylus brasilensis, the straight ventral margin of the proximal part of the deltopectoral crest, at first thought no greater precision was possible than a Pterodactyloidae incertae sedis
, but in 2000 narrowed it down to a Pteranodontoidea sensu Kellner. S. pricei according to Kellner belonged to a clade
descending from the last common ancestor of Istiodactylus
and the Anhangueridae. The same was in his analysis true for Araripesaurus
, a genus of which he had previously thought S. pricei was a junior synonym.
of 2.9–5.7 m (9.5–18.7 ft). De Buisonjé thought Santanadactylus brasilensis had a wingspan of six metres. It may have been adapted for gliding over flapping flight
.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of pterodactyloid pterosaur
Pterosaur
Pterosaurs were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic to the end of the Cretaceous Period . Pterosaurs are the earliest vertebrates known to have evolved powered flight...
from the Aptian
Aptian
The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous epoch or series and encompasses the time from 125.0 ± 1.0 Ma to 112.0 ± 1.0 Ma , approximately...
-age Romualdo Member of the Lower Cretaceous Santana Formation, of Barra do Jardim, Araripe Plateau, Ceará Province
Ceará
Ceará is one of the 27 states of Brazil, located in the northeastern part of the country, on the Atlantic coast. It is currently the 8th largest Brazilian State by population and the 17th by area. It is also one of the main touristic destinations in Brazil. The state capital is the city of...
, Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...
. Four species have been named, but today it is doubted they are part of the same genus. It was a rather large pterosaur.
History and Species
The genus was named in 1980 by the Dutch paleontologist Paul de Buisonjé. The type speciesType species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...
is S. brasilensis, the specific name referring to Brazil. It is based on holotype
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...
UvA M 4894 (Geological Institute of the University of Amsterdam), an upper part of the right humerus
Humerus
The humerus is a long bone in the arm or forelimb that runs from the shoulder to the elbow....
and a right scapula
Scapula
In anatomy, the scapula , omo, or shoulder blade, is the bone that connects the humerus with the clavicle ....
coracoid; other remains were also included in the genus: this paratype
Paratype
Paratype is a technical term used in the scientific naming of species and other taxa of organisms. The exact meaning of the term paratype when it is used in zoology is not the same as the meaning when it is used in botany...
consisted of two cervical
Neck
The neck is the part of the body, on many terrestrial or secondarily aquatic vertebrates, that distinguishes the head from the torso or trunk. The adjective signifying "of the neck" is cervical .-Boner anatomy: The cervical spine:The cervical portion of the human spine comprises seven boney...
vertebrae from a different individual, referred because they were found in the same lot of 25 chalk nodules bought from collectors. Additional remains, including a notarium
Notarium
Notarium is a term used for 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 adjacent vertebrae and ribs, while in some pterosaurs the notarium...
(fused vertebrae supporting the shoulder
Shoulder
The human shoulder is made up of three bones: the clavicle , the scapula , and the humerus as well as associated muscles, ligaments and tendons. The articulations between the bones of the shoulder make up the shoulder joints. The major joint of the shoulder is the glenohumeral joint, which...
) have been referred to it since then. The structure of the humerus suggests an ornithocheirid
Ornithocheiridae
Ornithocheiridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth.-Classification:Listing of genera after Unwin , except where noted.* Family Ornithocheiridae** ?Aetodactylus...
, but the long neck vertebrae argue against this.
In 1985, Peter Wellnhofer
Peter Wellnhofer
Peter Wellnhofer 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...
, a German paleontologist who has written numerous scientific publications on pterosaurs, named three additional species: S. araripensis, S. pricei, and S. spixi. S. araripensis, named after the Araripe Plateau, 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
Llewellyn Ivor Price was one of the first Brazilian paleontologists.-Biography:Price's work contributed not only to the development of Brazilian but also to global paleontology...
, was the smallest of the three species; it was based on BSP 1980 I 122, a left wing from the elbow
Elbow
The human elbow is the region surrounding the elbow-joint—the ginglymus or hinge joint in the middle of the arm. Three bones form the elbow joint: the humerus of the upper arm, and the paired radius and ulna of the forearm....
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 honouring Johann Baptist von Spix
Johann Baptist von Spix
Dr. Johann Baptist Ritter von Spix was a German naturalist.Spix was born in Höchstadt, Middle Franconia, as the seventh of eleven children. His boyhood home is the site of the Spix Museum , opened to the public in 2004...
.
Over the years, the species of this taxon have been reassessed. Chris Bennett suggested that S. brasilensis was a chimera
Chimera (paleontology)
In paleontology, a chimera is a fossil which was reconstructed with elements coming from more than a single species of animal. A now classic example of chimera is Protoavis.-List of paleontological chimeras:*Brontosaurus*Lametasaurus...
of a pteranodontid
Pteranodontidae
The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America.The family was named in 1876 by Othniel Charles Marsh....
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
Dsungaripteridae is a group of pterosaurs within the suborder Pterodactyloidea....
. Wellnhofer removed S. spixi from the genus as well, in 1991. In 1992, Alexander Kellner and Diogenes de Almeida Campos suggested that S. spixi was a tapejarid
Tapejaridae
Tapejaridae are a family of pterodactyloid pterosaurs from the early Cretaceous period. Members are currently known from Brazil and China, where the most primitive genera are found, indicating that the family has an Asian origin....
; David Unwin in 2003 thought that "Santanadactylus" spixi was a species of Tupuxuara
Tupuxuara
Tupuxuara is a genus of large, crested, toothless pterodactyloid pterosaur.The genus was in 1988 named and described by Alexander Kellner and Diogenes de Almeida Campos. The type species is Tupuxuara longicristatus. The genus name refers to a familiar spirit from the mythology of the Tupi...
. Kellner in 1990 renamed S. araripensis to Anhanguera
Anhangüera
Anhangüera originates from a word in the Tupi language which was the nickname given by Brazilian Indians to an explorer and "bandeirante", Bartolomeu Bueno da Silva .It may also refer to:* Anhanguera, Goiás, a municipality in the state of Goiás...
araripensis, followed by Wang and colleagues in 2008, though Veldmeijer (2003) included it in Coloborhynchus
Coloborhynchus
Coloborhynchus is a genus in the pterosaur family Ornithocheiridae, and is known from the Lower Cretaceous of England , and possibly the Aptian age of Brazil and Texas, depending on which species are included.-Description:The type specimen of Coloborhynchus is known only from a partial upper jaw...
. Part of the problem is the complicated taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...
of Santana Formation pterosaurs and their English
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...
contemporaries, involving numerous genera, such as Amblydectes, Anhanguera, Araripesaurus
Araripesaurus
Araripesaurus was a pterosaur, belonging to the Pterodactyloidea, from the famous Santana Formation of Brazil, dating from the Lower Cretaceous....
, Criorhynchus, Coloborhynchus, Lonchodectes
Lonchodectes
Lonchodectes was a genus of pterosaur from several formations dating to the Turonian of England, mostly in the area around Kent...
, Ornithocheirus
Ornithocheirus
Ornithocheirus was a pterosaur from the Cretaceous period of Europe and South America...
, and Tropeognathus.
Also the phylogeny is contentious. De Buisonjé first placed the genus into the Criorhynchidae, a concept that is no longer used. Wellnhofer considered it a member of the Ornithocheiridae
Ornithocheiridae
Ornithocheiridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth.-Classification:Listing of genera after Unwin , except where noted.* Family Ornithocheiridae** ?Aetodactylus...
. According to Bennett S. brasilensis belonged to Pteranodontidae
Pteranodontidae
The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America.The family was named in 1876 by Othniel Charles Marsh....
sensu Bennett. Kellner, concluding that he could find but a single autapomorphy
Autapomorphy
In cladistics, an autapomorphy is a distinctive anatomical feature, known as a derived trait, that is unique to a given terminal group. That is, it is found only in one member of a clade, but not found in any others or outgroup taxa, not even those most closely related to the group...
for Santanadactylus brasilensis, the straight ventral margin of the proximal part of the deltopectoral crest, at first thought no greater precision was possible than a Pterodactyloidae incertae sedis
Incertae sedis
, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...
, but in 2000 narrowed it down to a Pteranodontoidea sensu Kellner. S. pricei according to Kellner belonged to a clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
descending from the last common ancestor of Istiodactylus
Istiodactylus
Istiodactylus was a medium sized pterosaur from the Early Cretaceous period.-History and species:In 1887 Harry Govier Seeley named the remains of a fossil pelvis discovered on the Isle of Wight, Ornithodesmus cluniculus, thinking it was a bird. In 1901 he considered that it might have been a...
and the Anhangueridae. The same was in his analysis true for Araripesaurus
Araripesaurus
Araripesaurus was a pterosaur, belonging to the Pterodactyloidea, from the famous Santana Formation of Brazil, dating from the Lower Cretaceous....
, a genus of which he had previously thought S. pricei was a junior synonym.
Paleobiology
Santanadactylus is regarded as a large pterosaur, Wellnhofer for the various species indicating a wingspanWingspan
The wingspan of an airplane or a bird, is the distance from one wingtip to the other wingtip. For example, the Boeing 777 has a wingspan of about ; and a Wandering Albatross caught in 1965 had a wingspan of , the official record for a living bird.The term wingspan, more technically extent, is...
of 2.9–5.7 m (9.5–18.7 ft). De Buisonjé thought Santanadactylus brasilensis had a wingspan of six metres. It may have been adapted for gliding over flapping flight
Flight
Flight is the process by which an object moves either through an atmosphere or beyond it by generating lift or propulsive thrust, or aerostatically using buoyancy, or by simple ballistic movement....
.
External links
- Santanadactylus in The Pterosauria. Accessed 2007–02–09
- Re: Pterosaur Help, a posting from George OlshevskyGeorge OlshevskyGeorge Olshevsky is a freelance editor, writer, publisher, amateur paleontologist, and mathematician living in San Diego, California.Olshevsky maintains the comprehensive online Dinosaur Genera List...
on the Dinosaur Mailing List, which, although incomplete, should give some idea as to the complexity of the taxonomy here. Accessed 2007–02–09