Rhamphorhynchoidea
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The Rhamphorhynchoidea forms one of the two
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
s of
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 ...
s and represents an evolutionary grade of primitive members of flying
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...
s. This suborder is
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
unlike the
Pterodactyloidea 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 ...
, which arose from within the Rhamphorhynchoidea as opposed to a more distant common ancestor. Because it is not a completely natural grouping, Rhamphorhynchoidea is not used as a formal group in most scientific literature, though some pterosaur scientists continue to use it as an informal grouping in popular works, such as ''The Pterosaurs: From Deep Time'' by David Unwin, and in some formal studies. Rhamphorhynchoids were the first pterosaurs to have appeared, in the late
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest per ...
Period (
Norian The Norian is a division of the Triassic Period. It has the rank of an age ( geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic ...
age, about 210 million years agoButler, R.J., Barrett, P.M., and gower, D.J. (2009). "Postcranial skeletal pneumaticity and air-sacs in the earliest pterosaurs." ''Biology Letters'', 5(4): 557–560. ). Unlike their descendants, the pterodactyloids, most rhamphorhynchoids had teeth and long tails, and most species lacked a bony crest, though several are known to have crests formed from soft tissue like
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up Scale (anatomy), scales, hair, Nail ...
. They were generally small, with wingspans rarely exceeding 2.5 meters, though one specimen alluded to by
Alexander Stoyanow Alexander is a male given name. The most prominent bearer of the name is Alexander the Great, the king of the Ancient Greek kingdom of Macedonia (ancient kingdom), Macedonia who created one of the largest empires in ancient history. Variants li ...
would be among the largest pterosaurs of all time with a wingspan of 10 meters, comparable to the largest
azhdarchid Azhdarchidae (from the Persian word , , a dragon-like creature in Persian mythology) is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cre ...
s. However, this alleged giant Jurassic pterosaur specimen is not recorded anywhere outside the original ''
Time Time is the continued sequence of existence and event (philosophy), events that occurs in an apparently irreversible process, irreversible succession from the past, through the present, into the future. It is a component quantity of various me ...
'' article. Nearly all rhamphorhynchoids had become extinct by the end of the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
Period, though some anurognathids persisted to the early Cretaceous. The family
Wukongopteridae Wukongopteridae is a group of basal pterosaurs, found in China and the UK. It contains eight species in five genera, all dated to the Middle to Late Jurassic period, The Wukongopteridae were first named by Wang ''et al.'' in 2009, not yet givin ...
, which shows a mix of rhamphorhynchoid and pterodactyloid features, is known from the
Daohugou Beds The Haifanggou Formation (), also known as the Jiulongshan Formation (), is a fossil-bearing rock deposit located near Daohugou () village of Ningcheng County, in Inner Mongolia, northeastern China. The formation consists of coarse conglomerat ...
which are most commonly dated to the Jurassic, but a few studies give a Cretaceous date. Furthermore, remains of a non-pterodactyloid from the
Candeleros Formation The Candeleros Formation (formerly known as the Candeleros Member of the "Río Limay Formation") is a geologic formation that crops out in the Río Negro, Neuquén, and Mendoza provinces of northern Patagonia, Argentina. It is the oldest formati ...
extend the presence of basal pterosaurs into at least the early Late Cretaceous.Haluza, A. Y Apesteguía, S, Pterosaur remains (Archosauria, Ornithodira) from the early Late Cretaceous of “La Buitrera”, Río Negro, Argentina, Jornada; XXIII Jornadas Argentinas de Paleontología de Vertebrados; 2007


Classification


Taxonomy

Listing of families and superfamilies within the suborder Rhamphorhynchoidea, after Unwin 2006 unless otherwise noted. * Order Pterosauria ** Suborder Rhamphorhynchoidea * ***Clade Eopterosauria **** ''
Preondactylus ''Preondactylus'' is a genus of long-tailed pterosaurs from the Late Triassic (late Norian)Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an atlas. ''Zitteliana'', 61-107/ref> that ...
'' ****Clade Eudimorphodontimorpha *****''
Austriadactylus ''Austriadactylus'' is a genus of "rhamphorhynchoid" pterosaur. The fossil remains were unearthed in Late Triassic (middle Norian ageBarrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: ...
'' *****Clade
Eudimorphodontia Eudimorphodontidae is an extinct family of early pterosaurs from the Late Triassic (Late Carnian to Late Rhaetian age) of Europe.Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an ...
******''
Peteinosaurus ''Peteinosaurus'' ( ; meaning "winged lizard") was a prehistoric genus of pterosaur. It lived in the late Triassic period in the late Norian age (about 221 to 210 million years ago), and at a wingspan of around , was one of the smallest and earli ...
'' ******Family
Eudimorphodontidae Eudimorphodontidae is an extinct family of early pterosaurs from the Late Triassic (Late Carnian to Late Rhaetian age) of Europe.Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an a ...
*******Subfamily
Eudimorphodontinae Eudimorphodontidae is an extinct family of early pterosaurs from the Late Triassic (Late Carnian to Late Rhaetian age) of Europe.Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an a ...
********''
Eudimorphodon ''Eudimorphodon'' was a pterosaur that was discovered in 1973 by Mario Pandolfi in the town of Cene, Italy and described the same year by Rocco Zambelli. The nearly complete skeleton was retrieved from shale deposited during the Late Triassic (m ...
'' *******Subfamily
Raeticodactylinae Eudimorphodontidae is an extinct family of early pterosaurs from the Late Triassic (Late Carnian to Late Rhaetian age) of Europe.Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribution in time and space: an a ...
********''
Caviramus ''Caviramus'' is a genus of caviramid pterosaur from the Late Triassic (early Rhaetian-age) lower Kössen Formation of the Northern Calcareous Alps of Switzerland. The genus was in 2006 named by Nadia Fröbisch and Jörg Fröbisch. The type ...
'' ********''
Carniadactylus ''Carniadactylus'' is a genus of pterosaur which existed in Europe during the Late Triassic period (late Carnian or early Norian, about 228 million years agoBarrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). Pterosaur distribu ...
''? *** Family
Dimorphodontidae Dimorphodontidae (or dimorphodontids) is a group of early "rhamphorhynchoid" pterosaurs named after ''Dimorphodon'', that lived in the Late Triassic to Early Jurassic. While fossils that can be definitively referred to the group are rare, dimorph ...
*** Family
Anurognathidae Anurognathidae is a family of small, short-tailed pterosaurs that lived in Europe, Asia, and possibly North America during the Jurassic and Cretaceous periods. Five genera are known: '' Anurognathus'', from the Late Jurassic of Germany; ''Jeholop ...
*** Superfamily
Campylognathoidea Novialoidea (meaning "new wings") is an extinct clade of macronychopteran pterosaurs that lived from the latest Early Jurassic to the latest Late Cretaceous (early Toarcian to late Maastrichtian ageBarrett, P. M., Butler, R. J., Edwards, N. P., & ...
**** Family
Campylognathoididae Novialoidea (meaning "new wings") is an extinct clade of macronychopteran pterosaurs that lived from the latest Early Jurassic to the latest Late Cretaceous (early Toarcian to late Maastrichtian ageBarrett, P. M., Butler, R. J., Edwards, N. P ...
*** Family
Rhamphorhynchidae Rhamphorhynchidae is a group of early pterosaurs named after ''Rhamphorhynchus'', that lived in the Late Jurassic. The family Rhamphorhynchidae was named in 1870 by Harry Govier Seeley.Seeley, H.G. (1870). "The Orithosauria: An Elementary Study o ...
**** Subfamily
Rhamphorhynchinae Rhamphorhynchidae is a group of early pterosaurs named after ''Rhamphorhynchus'', that lived in the Late Jurassic. The family Rhamphorhynchidae was named in 1870 by Harry Govier Seeley.Seeley, H.G. (1870). "The Orithosauria: An Elementary Study o ...
**** Subfamily
Scaphognathinae Rhamphorhynchidae is a group of early pterosaurs named after ''Rhamphorhynchus'', that lived in the Late Jurassic. The family Rhamphorhynchidae was named in 1870 by Harry Govier Seeley.Seeley, H.G. (1870). "The Orithosauria: An Elementary Study o ...
* Rhamphorhynchoids of uncertain relationships (''
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, uncertain ...
'') ** '' Comodactylus'' ** '' Laopteryx'' ** ''
Parapsicephalus ''Parapsicephalus'' (meaning "beside arch head") is a genus of long-tailed rhamphorhynchid pterosaurs from the Lower Jurassic Whitby, Yorkshire, England. It contains a single species, ''P. purdoni'', named initially as a species of the related r ...
'' ** '' Rhamphinion''


References

* Unwin, D. M., (2003). "On the phylogeny and evolutionary history of pterosaurs." In Buffetaut, E. & Mazin, J.-M., eds. ''Evolution and Palaeobiology of Pterosaurs''. London: Geological Society of London, Special Publications 217, 2003, pp. 139–190. {{Taxonbar, from=Q1779482 Pterosaurs Paraphyletic groups