Klobiodon
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''Klobiodon'' is a genus of
rhamphorhynchid 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 ...
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
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
Taynton Limestone Formation The Taynton LimestoneWeishampel, David B; et al. (2004). "Dinosaur distribution (Middle Jurassic, Europe)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria, 2nd, Berkeley: University of California Press. Pp. 5 ...
of
Oxfordshire Oxfordshire is a ceremonial and non-metropolitan county in the north west of South East England. It is a mainly rural county, with its largest settlement being the city of Oxford. The county is a centre of research and development, primarily ...
, England.


Etymology

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 ...
of ''Klobiodon'' is ''Klobiodon rochei''. The generic name ''Klobiodon'' means "small cage tooth," from the Greek κλωβίον, ''klobion'', "little cage", and ὀδών, ''odon'', "tooth", in reference to the large anterior laniaries that appear to form a fish grab, while the specific name ''rochei'' honors the comic book artist Nick Roche for his anatomically correct designs inspired by dinosaurs.


History

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 ...
of ''Klobiodon rochei'', NHMUK PV OR 47991, was first mentioned by
George Robert Waterhouse George Robert Waterhouse (6 March 1810 – 21 January 1888) was an English natural history, naturalist. He was a keeper at the department of geology and later curator of the Zoological Society of London's museum. Early life George was born ...
(1878) as part of the collection of the
British Museum of Natural History The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum an ...
, where he mentions
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Owe ...
intended to name the specimen ''Pterodactylus raptor'' in an unpublished manuscript. It had been donated by
Robert Marsham Robert Marsham (27 January 17084 September 1797) was an English naturalist considered to be the founding father of phenology, the study of the effects of the seasons on plants and animals. Life He was admitted to Clare College, Cambridge in 172 ...
.
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...
(1888) referred the specimen to ''Rhamphorhynchus depressirostris'' (listed by him as '' Rhamphocephalus depressirostris'' and now known as ''
Dolichorhamphus ''Dolicorhamphus'' is an extinct genus of pterosaurs from the Middle Jurassic Taynton Limestone Formation and Fuller's Earth Formations of England. The genus contains two species, ''D. bucklandii'' and ''D. depressirostris''. Both species of ''D ...
depressirostris''). In their revision of pterosaur remains from the
Great Oolite Group The Great Oolite Group is a Middle Jurassic stratigraphic unit that outcrops in southern England. It consists of a complex set of marine deposits primarily mudstone and bioclastic ooidal and fine grained limestone, deposited in nearshore to shel ...
, Michael O'Sullivan and David Martill (2018) declared ''Rhamphorhynchus depressirostris'' a ''nomen dubium'' at ?Scaphognathinae indeterminatae, coining ''Klobiodon rochei'' for NHMUK PV OR 47991. A second specimen, OUM J.28410, was referred to the species but only provisionally because it lacks the, diagnostic, teeth.O’Sullivan, M. and Martill, D.M. 2018. Pterosauria of the Great Oolite Group (Bathonian, Middle Jurassic) of Oxfordshire and Gloucestershire, England. Acta Palaeontologica Polonica 63 (X): xxx–xxx. http://www.app.pan.pl/archive/published/app63/app004902018.pdf


Description

''Klobiodon'' is one of the largest pterosaurs known from the middle Jurassic. Its adult wingspan was estimated at two metres. The describing authors established a number of distinguishing traits. The lower jaw shows a unique combination of elongated front fang-like laniaries with short and robust teeth in a more backward position. The teeth more to the rear are at least 1.3 times longer than their tooth-sockets are wide. The rear laniary is at least 1.4 times and perhaps 2.4 times longer than the tooth to its rear (which could be an only partially erupted replacement tooth). The longest laniaries are at least 1.5 times taller than the vertical thickness of the front tooth-bearing lower jaw bone, the dentary, at its tallest point. The right lower jaw has a preserved length of fourteen centimetres. To the front, it gradually curves to below. A piece at the front has broken off. The original length of the jaw including this piece was estimated at eighteen centimetres. The missing piece was assumed by the authors to have borne one additional tooth. The fossil shows four teeth, which then would have been the second to fifth. The second and third teeth are elongated, recurved and inclined to the front. Such teeth are called laniaries. The fourth and fifth teeth are noticeably shorter but almost as wide, giving them a robust appearance. They are also straighter, especially at their rear edges, and perpendicular to the jaw.


Phylogeny

''Klobiodon'' was placed in the
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 ...
in 2018. At several places in the describing article, a position in the
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 ...
is suggested, but the authors eventually determine that it might also be member of the
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 ...
, concluding: "''Klobiodon rochei'' is therefore conservatively identified here as a member of the Rhamphorhynchidae rather than assigned to either subgroup". ''Klobiodon'' was found to be the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to the contemporaneous ''
Dolicorhamphus ''Dolicorhamphus'' is an extinct genus of pterosaurs from the Middle Jurassic Taynton Limestone Formation and Fuller's Earth Formations of England. The genus contains two species, ''D. bucklandii'' and ''D. depressirostris''. Both species of ''D ...
'' by Andres (2021).


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

{{Taxonbar, from=Q58403912 Jurassic pterosaurs of Europe Fossil taxa described in 2018 Monotypic prehistoric reptile genera Rhamphorhynchids