Banguela Oberlii
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''Banguela'' is 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 ...
of
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 ...
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
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 ...
. Only one species is known, ''Banguela oberlii''.


Discovery and naming

The Swiss collector Urs Oberli acquired a pterosaur jaw fragment from the
Chapada do Araripe 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 fl ...
. In 2005, this was described by André Jacques Veldmeijer and colleagues, and referred to ''
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 ...
sethi''.Veldmeijer, A.J., Signore, M. & Meijer, H.J.M., 2005, "Description of two pterosaur (Pterodactyloidea) mandibles from the lower Cretaceous Santana Formation, Brazil", ''DEINSEA'' 11: 67–86 In 2014 it was named and described by Jaime Headden and Hebert Bruno Nascimento Campos as a separate genus ''Banguela'', with 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 ...
''Banguela oberlii''. The generic name is a
Brazilian Portuguese Brazilian Portuguese (' ), also Portuguese of Brazil (', ) or South American Portuguese (') is the set of varieties of the Portuguese language native to Brazil and the most influential form of Portuguese worldwide. It is spoken by almost all of ...
word for "toothless one", especially used as an affectionate term for elderly women. 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 ...
honors Oberli. 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 ...
, NMSG SAO 251093, was probably found in 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 ...
, also known as the
Romualdo Member 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 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 stag ...
, dating 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 ...
. It consists of the
symphysis A symphysis (, pl. symphyses) is a fibrocartilaginous fusion between two bones. It is a type of cartilaginous joint, specifically a secondary cartilaginous joint. # A symphysis is an amphiarthrosis, a slightly movable joint. # A growing together ...
, fused front end, of the lower jaws. In 2018, a study placed the specimen in the subfamily
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 ...
, and formally named the species ''Thalassodromeus oberlii''.Rodrigo V. Pêgas, Fabiana R. Costa & Alexander W. A. Kellner, 2018, "New information on the osteology and a taxonomic revision of the genus ''Thalassodromeus'' (Pterodactyloidea, Tapejaridae, Thalassodrominae)", ''Journal of Vertebrate Paleontology'' Article: e1443273


Description

''Banguela'' has an estimated skull length of about and
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 over . The symphysis, with a preserved length of , curves upwards and has a relatively short depression at its upper rear end. The front upper edge of the symphysis is sharp. The front bottom edge is sharp too but lacks a true crest. There are no teeth or tooth sockets present in the fragment.


Phylogeny

In 2005, Veldmeijer had already noted similarities to ''
Dsungaripterus ''Dsungaripterus'' is a genus of dsungaripterid pterosaur with an average wingspan of . ''Dsungaripterus'' lived during the Early Cretaceous in what is now China, and its first fossil was found in the Junggar Basin. Description ''Dsungaripter ...
'', but considered the available data to be insufficient to draw any conclusions from this. In 2014, Headden & Campos placed ''Banguela'' in the family
Dsungaripteridae 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 ...
, in a basal position. The cladogram of their analysis is shown below: ''Banguela'' is unique among dsungaripterid pterosaurs due to a presumed total absence of teeth. Other pterosaur groups, such as
pteranodontids 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 ( ...
,
nyctosaurids Nyctosauridae (meaning "night lizards" or "bat lizards") is a Family (biology), family of specialized soaring pterosaurs of the late Cretaceous Period of North America, Africa, and possibly Europe. It was named in 1889 by Henry Alleyne Nicholson ...
and azhdarchoids, have also lost their teeth, indicating that toothloss might have independently occurred at least four times among pterosaurs. However, because dsungaripterids are occasionally recovered as derived azhdarchoids, it is possible that toothloss has occurred more often, if as an instance of
Dollo's Law Dollo's law of irreversibility (also known as Dollo's law and Dollo's principle), proposed in 1893 by Belgian paleontologist Louis Dollo states that, "an organism never returns exactly to a former state, even if it finds itself placed in condition ...
azhdarchoids should be originally toothless. If there was a large number of cases, ''Banguela'' suggests how it developed in most of these: the development of horned rhamphothecae in the jawtips, with progressive tooth rarification until they cease to be useful. It is worth to note that dsungaripteroids have some of the most specialized teeth of all
sauropsids Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early synap ...
, so ''Banguelas toothlessness must indicate some degree of divergent specialization. In 2018, a study suggested that ''Banguela'' is a species of ''Thalassodromeus'' (''T. oberlii''), and thus it is assigned to the Thalassodrominae, a subfamily within the clade
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 ...
.


See also

*
List of pterosaur genera This list of pterosaurs is a comprehensive listing of all genera that have ever been included in the order Pterosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considered inval ...
*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs w ...


References

{{Portal bar, Paleontology, Cretaceous, Brazil Azhdarchoids Early Cretaceous pterosaurs of South America Cretaceous Brazil Fossils of Brazil Fossil taxa described in 2014