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''Caulkicephalus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
anhanguerid Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a late ...
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 ...
from the
Isle of Wight The Isle of Wight ( ) is a Counties of England, county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the List of islands of England#Largest islands, largest and List of islands of England#Mo ...
off the coast of
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...
. It lived during the Early Cretaceous period, about 130 million years ago.


Discovery and naming

Between 1995 and 2003, bone fragments of an unknown pterosaur were found at the
Yaverland Yaverland is a village on the Isle of Wight, just north of Sandown on Sandown Bay. It has about 200 houses. About of a mile away from the village is the Yaverland Manor and Church. Holotype fossils have been discovered here of ''Yaverlan ...
locality near
Sandown Sandown is a seaside resort and civil parish on the south-east coast of the Isle of Wight, United Kingdom with the resort of Shanklin to the south and the settlement of Lake in between. Together with Shanklin, Sandown forms a built-up area of ...
. The discoveries were made in or from a brown clay layer from the
Wessex Formation The Wessex Formation is a fossil-rich English geological formation that dates from the Berriasian to Barremian stages (about 145–125 million years ago) of the Early Cretaceous. It forms part of the Wealden Group and underlies the younger Vect ...
of the Wealden Group, stemming from the Early Cretaceous (
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is precede ...
age, about 130 million years ago). In 2005 a new genus was named and described by Lorna Steel, David Martill,
David Unwin David (; , "beloved one") (traditional spelling), , ''Dāwūd''; grc-koi, Δαυΐδ, Dauíd; la, Davidus, David; gez , ዳዊት, ''Dawit''; xcl, Դաւիթ, ''Dawitʿ''; cu, Давíдъ, ''Davidŭ''; possibly meaning "beloved one". w ...
and John Winch. 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 specime ...
is ''Caulkicephalus trimicrodon''. The genus name is a translation of "Caulkhead", a traditional nickname for Isle of Wight residents, partially derived from Greek ''kephale'', "head". The specific name, ''trimicrodon'', means "three small teeth", in reference to the dentition.


Description

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 sever ...
is IWCMS 2002.189.1, 2, 4: three pieces, more or less contiguous, of the front part of a snout. As
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...
s have been referred: IWCMS 2002.189.3, a partial posterior skull roof; IWCMS 2003.2, a left quadrate; IWCMS 2003.4, a possible partial
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
; ICWMS 2002.237, a 44 millimeter (1.7 in) long fragment of the first phalanx of the wing finger; IWCMS 2002.234.1-4, four contiguous fragments of a first phalanx, together measuring long; IWCMS 2002.233, a possible distal end, long, of a second phalanx; IWCMS 2002.236, a fragment of the shaft of possibly the fourth phalanx; and IWCMS 2003.3, a probable fragment of a hindlimb bone. The fossils have only been slightly compressed. The snout fragments have a combined length of . On the snout top the base of a crest is visible, not quite reaching its rounded tip. The teeth have, apart from some replacement teeth present deep in the jaw, been lost but their number, orientation and size can be inferred from the tooth sockets, which however are partly missing at the right side. These are oval and slightly elevated above the jaw bone. The first two tooth pairs were pointed somewhat to the front; the teeth more to the back pointed more sideways; the most posterior preserved stood perpendicular to the jaw. The teeth increased in size until the third pair which was the largest. The fourth pair was equal to the first but the fifth, sixth and seventh pairs were markedly smaller, less than half in size; it is this feature which is recalled by the specific name. Pairs eight, nine and ten again equalled the first. After a narrow hiatus between the second and third snout fragment four tooth sockets are present at each side of the latter, but these are not placed in opposite pairs. The number of teeth in the upper jaw thus seems to have been at least fourteen. The smaller sized teeth were placed in a constriction of the snout, which thus had a broader end with larger teeth, a so-called "prey grab", usually interpreted as an adaptation to catch slippery prey such as fish. The posterior skull fragment, a braincase which is rather damaged, shows on its top the base of a parietal crest, probably pointing towards the back. It seems to have been separate from the snout crest. The layer the fossils were found in, does not consist of marine sediments, but contains land plant debris; this is seen as an indication of a more terrestrial habitat. David Martill estimated ''Caulkicephalus'' had a
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 o ...
of around 5 meters (16.5 ft).


Classification

''Caulkicephalus'' was by the describers assigned to the
Ornithocheiridae Ornithocheiridae (or ornithocheirids, meaning "bird hands") is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the Early to Late Cr ...
in view of the narrowing in the middle of the snout. The snout crest was seen as an indication it belonged to the more general
Ornithocheiroidea Ornithocheiroidea (or ornithocheiroids) is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods (Valanginian to Maastrichtian stages), with fo ...
''sensu'' Unwin, whereas the parietal crest was suggested to have been a
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
, a shared new feature, of the more narrow group of the
Euornithocheira Pteranodontoidea (or pteranodontoids, from Greek meaning "toothless wings") is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous (early Valanginian to late Maastrichtian stages) of Asia, Africa, Europe, North Americ ...
. Unique characters of the species itself, its
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
, are the details of its dentition, the downward and backward running suture between the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
and
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. T ...
, and the fact the median ridge of the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly separ ...
begins (or ends) at the ninth tooth pair. In 2019 however, several studies have reassigned ''Caulkicephalus'' to the family
Anhangueridae Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a later ...
, specifically to the subfamily Anhanguerinae, sister taxon to both ''
Guidraco ''Guidraco'' (Chin. ''gui'' (鬼) "malicious ghost" + Lat. ''draco'' "dragon") is an extinct genus of toothed pterodactyloid pterosaur known from the Early Cretaceous of Liaoning Province, northeast China. According to many recent studies, ''Gu ...
'' and ''
Ludodactylus ''Ludodactylus'' (meaning "play finger") is a genus of anhanguerid pterodactyloid pterosaur from the Early Cretaceous period (Aptian stage) of what is now the Crato Formation of the Araripe Basin in Ceará, Brazil. The type and only species i ...
''.Borja Holgado, Rodrigo V. Pêgas, José Ignacio Canudo, Josep Fortuny, Taissa Rodrigues, Julio Company & Alexander W.A. Kellner, 2019, "On a new crested pterodactyloid from the Early Cretaceous of the Iberian Peninsula and the radiation of the clade Anhangueria", ''Scientific Reports'' 9: 4940 Zhou X., Pêgas R.V., Leal M.E.C. & Bonde N. 2019
"''Nurhachius luei'', a new istiodactylid pterosaur (Pterosauria, Pterodactyloidea) from the Early Cretaceous Jiufotang Formation of Chaoyang City, Liaoning Province (China) and comments on the Istiodactylidae."
''PeerJ'' 7:e7688
The cladogram below is a topology recovered by Rodrigo Pêgas and colleagues in 2019, showing the position of ''Caulkicephalus'' within the Anhangueridae.Rodrigo V. Pêgas, Borja Holgado & Maria Eduarda C. Leal (2019) On ''Targaryendraco wiedenrothi'' gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids, Historical Biology,


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 ...
* Pterosaur size


References


External links


Description, references, and images of ''Caulkicephalus''.
{{Portal bar, Paleontology, Cretaceous, United Kingdom Pteranodontoids Early Cretaceous pterosaurs of Europe Fossil taxa described in 2005