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{{short description|Genus of azhdarchoid pterosaur from the Early Cretaceous}}
{{Italic title}}
{{Speciesbox
{{Taxobox
⚫
|fossil_range={{fossil range|Albian}}
⚫
|name = ''Banguela''
⚫
|fossil_range = [[Albian]]
|image=Banguela oberlii.png
|image=Banguela oberlii.png
|image_caption=Illustration of the holotype jaw fragment, with cross-section
|image_caption=Illustration of the holotype jaw fragment, with cross-section
⚫
|genus=Banguela
|regnum = [[Animal]]ia
⚫
|parent_authority=Headden & Campos, 2015
|phylum = [[Chordate|Chordata]]
|species=oberlii
|classis = [[Reptile|Sauropsida]]
⚫
|authority=Headden & Campos, 2015
|ordo = [[Pterosaur]]ia
|display_parents = 2
|subordo = [[Pterodactyloidea]]
⚫
|synonyms=*''[[Thalassodromeus]] oberlii''? <br />{{small|Pêgas, Costa & Kellner, 2018}}
⚫
|familia = [[Dsungaripteridae]]
⚫
|genus = '''''Banguela'''''
⚫
|genus_authority = Headden & Campos 2014
|binomial = ''Banguela oberlii''
⚫
|binomial_authority = Headden & Campos, 2013
| synonyms =
⚫
* ''Thalassodromeus oberlii''? {{small|(Headden & Campos, 2013)}}
}}
}}


'''''Banguela oberlii''''' is a [[Dsungaripteridae|dsungaripterid]] [[pterosaur]] from the [[Early Cretaceous]] of [[Brazil]].<ref name="Banguela">{{Cite journal|author=Jaime A. Headden and Hebert B.N. Campos |year=2014 |title=An unusual edentulous pterosaur from the Early Cretaceous Romualdo Formation of Brazil |journal=Historical Biology: An International Journal of Paleobiology |volume=in press |issue= |pages= |doi=10.1080/08912963.2014.904302 }}</ref>
'''''Banguela''''' is a [[genus]] of [[azhdarchoid]] [[pterosaur]] from the [[Early Cretaceous]] period ([[Albian]] stage) of what is now [[Brazil]]. Only one species is known, '''''Banguela oberlii'''''.<ref name="Banguela">{{Cite journal|author=Jaime A. Headden and Hebert B.N. Campos |year=2015 |title=An unusual edentulous pterosaur from the Early Cretaceous Romualdo Formation of Brazil |journal=Historical Biology: An International Journal of Paleobiology |volume=27 |issue=7 |pages=815–826 |doi=10.1080/08912963.2014.904302 |s2cid=129306469 }}</ref>


== Discovery and naming ==
==Discovery and naming==
⚫
The Swiss collector [[Urs Oberli]] acquired a pterosaur jaw fragment from the [[Chapada do Araripe]]. In 2005, this was described by [[André Jacques Veldmeijer]] and colleagues, and referred to ''[[Thalassodromeus]] sethi''.<ref name="Veldmeijer2005">{{cite journal | last1 = Veldmeijer | first1 = A.J. | last2 = Signore | first2 = M. | last3 = Meijer | first3 = H.J.M. | date = 2005 | title = Description of two pterosaur (Pterodactyloidea) mandibles from the upper Cretaceous Santana Formation, Brazil | journal = Deinsea | volume = 11 | pages = 67–86 }}</ref>


⚫
In 2015, it was named and described by [[Jaime Headden]] and [[Hebert Bruno Nascimento Campos]] as a separate genus ''Banguela'', with the [[type species]] ''Banguela oberlii''. The [[genus|generic name]] is a [[Brazilian Portuguese]] word for "toothless one", especially used as an affectionate term for elderly women. The [[specific name (zoology)|specific name]] honors Oberli.<ref name="Banguela"/>
⚫
The Swiss collector [[Urs Oberli]] acquired a pterosaur jaw fragment from the [[Chapada do Araripe]]. In 2005, this was described by [[André Jacques Veldmeijer]] e.a. and referred to ''[[Thalassodromeus]] sethi''.<ref name="Veldmeijer2005">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</ref>

⚫
In 2014 it was named and described by [[Jaime Headden]] and [[Hebert Bruno Nascimento Campos]] as a separate genus ''Banguela'', with the [[type species]] ''Banguela oberlii''. The genus name is a [[Brazilian Portuguese]] word for "toothless one", especially used as an affectionate term for elderly women. The [[specific name (zoology)|specific name]] honours Oberli.<ref name="Banguela"/>


The [[holotype]], '''NMSG SAO 251093''', was probably found in the [[Romualdo Formation]], also known as the [[Romualdo Member]] of the [[Santana Formation]], dating from the [[Albian]]. It consists of the [[symphysis]], fused front end, of the lower jaws.<ref name="Banguela"/>
The [[holotype]], '''NMSG SAO 251093''', was probably found in the [[Romualdo Formation]], also known as the [[Romualdo Member]] of the [[Santana Formation]], dating from the [[Albian]]. It consists of the [[symphysis]], fused front end, of the lower jaws.<ref name="Banguela"/>

In 2018, a study placed the specimen in the Thalassodrominae and formally named the species ''Thalassodromeus oberlii''.<ref name="Pêgas2018">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</ref>


==Description==
==Description==
[[File:Banguela skull.png|thumb|left|The specimen reconstructed as a [[dsungaripterid]] by Jaime Headden]]
[[File:Banguela skull.png|thumb|The specimen reconstructed as a [[dsungaripterid]] by Jaime Headden]]
''Banguela'' has an estimated skull length of about two feet and [[wingspan]] of over twelve feet. The symphysis, with a preserved length of 273 millimetres, 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.<ref name="Banguela"/>
''Banguela'' has an estimated skull length of about {{convert|2|ft|cm|order=flip|sp=us}} and [[wingspan]] of over {{convert|12|ft|m|order=flip|sp=us}}. The symphysis, with a preserved length of {{convert|273|mm|in|sp=us}}, 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.<ref name="Banguela"/>


==Phylogeny==
==Phylogeny==
Veldmeijer had already in 2005 noted similarities to ''[[Dsungaripterus]]'', but considered the available data to be insufficient to draw any conclusions from this.<ref name="Veldmeijer2005"/> In 2014, Headden & Campos placed ''Banguela'' in the [[Dsungaripteridae]], in a basal position. ''Banguela'' is unique among dsungaripterid pterosaurs due to a presumed total absence of teeth. Other pterosaur groups, such as [[Pteranodontidae|pteranodontids]], [[Nyctosauridae|nyctosaurids]] and [[Azhdarchoidea|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]] azhdarchoids should be originally toothless.<ref name="Banguela"/><ref name=LoneStarPterosaurs>{{Cite journal | last1 = Andres | first1 = B. | last2 = Myers | first2 = T. S. | doi = 10.1017/S1755691013000303 | title = Lone Star Pterosaurs | journal = Earth and Environmental Science Transactions of the Royal Society of Edinburgh | pages = 1 | year = 2013 | pmid = | pmc = }}</ref> 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.<ref name="Banguela"/>
In 2005, Veldmeijer had already noted similarities to ''[[Dsungaripterus]]'', but considered the available data to be insufficient to draw any conclusions from this.<ref name="Veldmeijer2005"/> In 2015, Headden & Campos placed ''Banguela'' in the family [[Dsungaripteridae]], in a basal position. The cladogram of their analysis is shown below:<ref name="Banguela"/>


{{clade| style=font-size:100%;line-height:100%
It is worth to note that dsungaripteroids have some of the most specialised teeth of all [[Sauropsida|sauropsids]],<ref>{{Cite book| title = Pterosaurs: Natural History, Evolution, Anatomy|author=Wilton, Mark P. |isbn=0691150613|year=2013|publisher=Princeton University Press }}</ref> so ''Banguela'''s toothlessness must indicate some degree of divergent specialisation.
|label1=[[Azhdarchoidea]]
|1={{clade
⚫
|label1=[[Dsungaripteridae]]
|1={{clade
⚫
|1='''''Banguela oberlii'''''
|2={{clade
|1=''[[Dsungaripterus weii]]''
|2=''[[Noripterus complicidens]]''
}}
}}
|label2=[[Neoazhdarchia]]
|2={{clade
|1={{clade
|1=[[Chaoyangopteridae]]
|2=[[Azhdarchidae]]
}}
|label2=[[Tapejaroidea]]
|2={{clade
|1=[[Thalassodromidae]]
|2=[[Tapejaridae]]
}} }} }} }}


If ''Banguela'' were a dsungaripterid pterosaur, it would have been unique among the group due to the presumed total absence of teeth. Other pterosaur groups, such as [[Pteranodontidae|pteranodontids]], [[Nyctosauridae|nyctosaurids]] and [[Azhdarchoidea|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]] azhdarchoids should be originally toothless.<ref name="Banguela"/><ref name=LoneStarPterosaurs>{{Cite journal |last1=Andres |first1=B. |last2=Myers |first2=T. S. |doi=10.1017/S1755691013000303 |title=Lone Star Pterosaurs |journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh |pages=383–398 |year=2013 |volume=103 |issue=3–4 |s2cid=84617119 }}</ref> 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.<ref name="Banguela"/> Since [[dsungaripteroid]]s have some of the most specialized teeth of all [[Sauropsida|sauropsids]],<ref>{{Cite book|title=Pterosaurs: Natural History, Evolution, Anatomy|author=Wilton, Mark P. |isbn=978-0691150611|year=2013|publisher=Princeton University Press }}</ref> the toothlessness of ''Banguela'' could indicate some degree of divergent specialization.
The 2018 placed the species in the [[Thalassodrominae]].<ref name="Pêgas2018"/>

However, its identity as a dsungaripterid was disputed by other researchers, since no unambiguous traits of dsungaripterids were found in the type specimen.<ref>{{cite journal | last1 = Pêgas | first1 = R.V. | last2 = Leal | first2 = M.E.d. | last3 = Kellner | first3 = A.W.A. | title = A Basal Tapejarine (Pterosauria; Pterodactyloidea; Tapejaridae) from the Crato Formation, Early Cretaceous of Brazil. | journal = PLOS ONE | date = 2016 | volume = 11 | issue = 9 | page = e0162692 | doi = 10.1371/journal.pone.0162692 | pmid=27655346 | pmc=5031394| bibcode = 2016PLoSO..1162692P | doi-access = free }}</ref><ref name="Pêgas2018"/><ref name=M2020/> In 2018, a study suggested that ''Banguela'' was a species of ''[[Thalassodromeus]]'' (''T. oberlii''), and thus it is assigned to the Thalassodrominae, a subfamily within the clade [[Tapejaridae]].<ref name="Pêgas2018">{{cite journal |last1=Pêgas |first1=R. V. |last2=Costa |first2=F. R. |last3=Kellner |first3=A. W. A. |title=New Information on the osteology and a taxonomic revision of The genus ''Thalassodromeus'' (Pterodactyloidea, Tapejaridae, Thalassodrominae) |journal=Journal of Vertebrate Paleontology |date=2018 |volume=38 |issue=2 |pages=e1443273 |doi=10.1080/02724634.2018.1443273|bibcode=2018JVPal..38E3273P |s2cid=90477315 }}</ref> In 2020, McPhee and colleagues considered ''Banguela'' as a valid genus, and claimed that it belongs to the [[Chaoyangopteridae]].<ref name=M2020>{{cite journal|last1=McPhee|first1=James|last2=Ibrahim|first2=Nizar|author-link2=Nizar Ibrahim|last3=Kao|first3=Alex|last4=Unwin|first4=David M.|last5=Smith|first5=Roy|last6=Martill|first6=David M.|year=2020|title=A new ?chaoyangopterid (Pterosauria: Pterodactyloidea) from the Cretaceous Kem Kem beds of Southern Morocco|journal=Cretaceous Research|volume=110|pages=Article 104410|doi=10.1016/j.cretres.2020.104410|s2cid=213739173 |url=https://researchportal.port.ac.uk/portal/en/publications/a-new-chaoyangopterid-pterosauria-pterodactyloidea-from-the-cretaceous-kem-kem-beds-of-southern-morocco(78a38938-aa6d-4802-973e-d646e1e6d594).html }}</ref>


==See also==
==See also==
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==References==
==References==
{{Reflist}}
{{Reflist}}
{{Pterosauria|Az.}}

{{Taxonbar|from=Q16740190}}
{{Taxonbar|from=Q16740190}}
{{Portal bar|Paleontology|Brazil}}


[[Category:Dsungaripteroids]]
[[Category:Azhdarchoids]]
[[Category:Late Cretaceous pterosaurs of South America]]
[[Category:Early Cretaceous pterosaurs of South America]]
[[Category:Cretaceous Brazil]]
[[Category:Cretaceous Brazil]]
[[Category:Fossils of Brazil]]
[[Category:Fossils of Brazil]]

Latest revision as of 20:09, 2 December 2024

Banguela
Temporal range: Albian
Illustration of the holotype jaw fragment, with cross-section
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order: †Pterosauria
Suborder: †Pterodactyloidea
Clade: †Azhdarchoidea
Genus: †Banguela
Headden & Campos, 2015
Species:
†B. oberlii
Binomial name
†Banguela oberlii
Headden & Campos, 2015
Synonyms

Banguela is a genus of azhdarchoid pterosaur from the Early Cretaceous period (Albian stage) of what is now Brazil. Only one species is known, Banguela oberlii.[1]

Discovery and naming

[edit]

The Swiss collector Urs Oberli acquired a pterosaur jaw fragment from the Chapada do Araripe. In 2005, this was described by André Jacques Veldmeijer and colleagues, and referred to Thalassodromeus sethi.[2]

In 2015, it was named and described by Jaime Headden and Hebert Bruno Nascimento Campos as a separate genus Banguela, with the type species Banguela oberlii. The generic name is a Brazilian Portuguese word for "toothless one", especially used as an affectionate term for elderly women. The specific name honors Oberli.[1]

The holotype, NMSG SAO 251093, was probably found in the Romualdo Formation, also known as the Romualdo Member of the Santana Formation, dating from the Albian. It consists of the symphysis, fused front end, of the lower jaws.[1]

Description

[edit]
The specimen reconstructed as a dsungaripterid by Jaime Headden

Banguela has an estimated skull length of about 61 centimeters (2 ft) and wingspan of over 3.7 meters (12 ft). The symphysis, with a preserved length of 273 millimeters (10.7 in), 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.[1]

Phylogeny

[edit]

In 2005, Veldmeijer had already noted similarities to Dsungaripterus, but considered the available data to be insufficient to draw any conclusions from this.[2] In 2015, Headden & Campos placed Banguela in the family Dsungaripteridae, in a basal position. The cladogram of their analysis is shown below:[1]

Azhdarchoidea

If Banguela were a dsungaripterid pterosaur, it would have been unique among the group due to the presumed total absence of teeth. Other pterosaur groups, such as pteranodontids, nyctosaurids 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 azhdarchoids should be originally toothless.[1][3] 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.[1] Since dsungaripteroids have some of the most specialized teeth of all sauropsids,[4] the toothlessness of Banguela could indicate some degree of divergent specialization.

However, its identity as a dsungaripterid was disputed by other researchers, since no unambiguous traits of dsungaripterids were found in the type specimen.[5][6][7] In 2018, a study suggested that Banguela was a species of Thalassodromeus (T. oberlii), and thus it is assigned to the Thalassodrominae, a subfamily within the clade Tapejaridae.[6] In 2020, McPhee and colleagues considered Banguela as a valid genus, and claimed that it belongs to the Chaoyangopteridae.[7]

See also

[edit]

References

[edit]
  1. ^ a b c d e f g Jaime A. Headden and Hebert B.N. Campos (2015). "An unusual edentulous pterosaur from the Early Cretaceous Romualdo Formation of Brazil". Historical Biology: An International Journal of Paleobiology. 27 (7): 815–826. doi:10.1080/08912963.2014.904302. S2CID 129306469.
  2. ^ a b Veldmeijer, A.J.; Signore, M.; Meijer, H.J.M. (2005). "Description of two pterosaur (Pterodactyloidea) mandibles from the upper Cretaceous Santana Formation, Brazil". Deinsea. 11: 67–86.
  3. ^ Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103 (3–4): 383–398. doi:10.1017/S1755691013000303. S2CID 84617119.
  4. ^ Wilton, Mark P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. ISBN 978-0691150611.
  5. ^ Pêgas, R.V.; Leal, M.E.d.; Kellner, A.W.A. (2016). "A Basal Tapejarine (Pterosauria; Pterodactyloidea; Tapejaridae) from the Crato Formation, Early Cretaceous of Brazil". PLOS ONE. 11 (9): e0162692. Bibcode:2016PLoSO..1162692P. doi:10.1371/journal.pone.0162692. PMC 5031394. PMID 27655346.
  6. ^ a b Pêgas, R. V.; Costa, F. R.; Kellner, A. W. A. (2018). "New Information on the osteology and a taxonomic revision of The genus Thalassodromeus (Pterodactyloidea, Tapejaridae, Thalassodrominae)". Journal of Vertebrate Paleontology. 38 (2): e1443273. Bibcode:2018JVPal..38E3273P. doi:10.1080/02724634.2018.1443273. S2CID 90477315.
  7. ^ a b McPhee, James; Ibrahim, Nizar; Kao, Alex; Unwin, David M.; Smith, Roy; Martill, David M. (2020). "A new ?chaoyangopterid (Pterosauria: Pterodactyloidea) from the Cretaceous Kem Kem beds of Southern Morocco". Cretaceous Research. 110: Article 104410. doi:10.1016/j.cretres.2020.104410. S2CID 213739173.
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