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''Camposipterus'' 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
pterodactyloid 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 ...
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 ...
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 ...
. Fossil remains of ''Camposipterus'' dated back to 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 ...
, about 112 million years ago.


Discovery and naming

In 1869,
Harry Govier Seeley Harry Govier Seeley (18 February 1839 – 8 January 1909) was a British paleontologist. Early life Seeley was born in London on 18 February 1839, the second son of Richard Hovill Seeley, a goldsmith, and his second wife Mary Govier. When his fat ...
, based on a fossil found at
Haslingfield Haslingfield is a village and civil parish in South Cambridgeshire, England. The village is about six miles south-west of Cambridge, between Harston, Barton and Barrington. The population in the 2001 census was 1,550 people living in 6 ...
,
Cambridgeshire Cambridgeshire (abbreviated Cambs.) is a Counties of England, county in the East of England, bordering Lincolnshire to the north, Norfolk to the north-east, Suffolk to the east, Essex and Hertfordshire to the south, and Bedfordshire and North ...
, named ''Ptenodactylus nasutus'',Seeley, H.G., 1869, ''Index to the fossil remains of Aves, Ornithosauria, and Reptilia, from the Secondary System of Strata arranged in the Woodwardian Museum of the University of Cambridge''. Deighton, Bell and Co., Cambridge, xxiii + 143 pp at the same time disclaiming the name which makes it invalid by modern standards. In 1870, Seeley had realized that the generic name ''Ptenodactylus'' had been preoccupied, so he renamed the species into ''
Ornithocheirus ''Ornithocheirus'' (from Ancient Greek "ὄρνις", meaning ''bird'', and "χεῖρ", meaning ''hand'') is a pterosaur genus known from fragmentary fossil remains uncovered from sediments in the UK and possibly Morocco. Several species have ...
nasutus''.Seeley, H.G., 1870, ''The Ornithosauria: an elementary study of the bones of pterodactyls, made from fossil remains found in the Cambridge Upper Greensand, and arranged in the Woodwardian Museum of the University of Cambridge''. Deighton, Bell, and Co., Cambridge, xii + 135 pp The specific name means "with a long nose" in
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
. In 2001,
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 ...
made this species a
junior subjective synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
of ''
Anhanguera Anhanguera may refer to: People * Bartolomeu Bueno da Silva (1672–1740), a bandeirante Places in Brazil * Anhanguera, Goiás, a municipality in the state of Goiás * Anhanguera (district of São Paulo), a district in São Paulo * Parque Anhan ...
fittoni''. However, in 2013, Taissa Rodrigues and Alexander Wilhelm Armin Kellner concluded firstly that ''Pterodactylus fittoni'' was not a part of the genus ''Anhanguera'' and secondly that ''Ornithocheirus nasutus'' was not identical to it regardless. They decided to name a separate genus for the species: ''Camposipterus''. The generic name combines that of the Brazilian paleontologist Diogenes de Almeida Campos with a Latinized Greek πτερόν, pteron, "wing". The resulting new combination name, the ''
combinatio nova ''Combinatio nova'', abbreviated ''comb. nov.'' (sometimes ''n. comb.''), is Latin for "new combination". It is used in taxonomic biology literature when a new name is introduced based on a pre-existing name. The term should not to be confused wi ...
'', which is ''Camposipterus nasutus'', while 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 ...
remains as ''Ornithocheirus nasutus''. 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 ...
, CAMSM B 54556, had been found in a layer of the
Cambridge Greensand The Cambridge Greensand is a geological unit in England whose strata are earliest Cenomanian in age. It lies above the erosive contact between the Gault Formation and the Chalk Group in the vicinity of Cambridgeshire, and technically forms the low ...
dating from the
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
but probably containing reworked fossils from the older
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 front part of a snout. Rodrigues & Kellner in 2013 moved two more species to the genus. They renamed ''Pterodactylus sedgwickii'' into ''Camposipterus''(?) ''sedgwickii'' and ''Ornithocheirus colorhinus'' into ''Camposipterus''(?) ''colorhinus''. The question marks indicate the uncertainty of the authors about the correctness of the move. ''Pterodactylus sedgwickii'' had in 1859 been named by
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 ...
based on specimen CAMSM B54422, the front part of a snout from the Cambridge Greensand. Its specific name honors
Adam Sedgwick Adam Sedgwick (; 22 March 1785 – 27 January 1873) was a British geologist and Anglican priest, one of the founders of modern geology. He proposed the Cambrian and Devonian period of the geological timescale. Based on work which he did on W ...
. It was in 1869 renamed by Seeley into a ''Ptenodactylus sedgwickii'', and in 1870 into a ''Ornithocheirus sedgwickii''. In 1874, Owen again renamed it into ''
Coloborhynchus ''Coloborhynchus'' is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. ''Coloborhynchus'' is known from the Lower Cretaceous of Engla ...
sedgwickii''.Owen, R., 1874, ''Monograph on the fossil Reptilia of the Mesozoic Formations''. Palaeontographical Society, London, 14 pp Owen in 1859 also referred a front of the lower jaws, specimen CAMSM B54421. However, this piece is not of the same individual as the holotype and there is no proof for any connection with ''Pterodactylus sedgwickii''. In 1869, Seeley also named a ''Ptenodactylus colorhinus'', in 1870 an ''Ornithocheirus colorhinus'' based on the
syntype In biological nomenclature, a syntype is any one of two or more biological types that is listed in a description of a taxon where no holotype was designated. Precise definitions of this and related terms for types have been established as part of ...
s CAMSM B54431 and CAMSM B54432, both front snouts from the Cambridge Greensand. Its specific name means "with a docked nose" from the Greek κόλος, ''kolos'', "docked", and ῥίς. ''rhis'', "nose". In 2001, Unwin considered this species to be a junior synonym of ''Anhanguera cuvieri''. However, this was not accepted by Rodrigues & Kellner, who, rejecting any identity between the Brazilian and English pterosaur material, named a separate ''
Cimoliopterus ''Cimoliopterus'' is a genus of pterosaur that lived during the Late Cretaceous in what is now England and the United States. The first known specimen, consisting of the front part of a snout including part of a crest, was discovered in the G ...
cuvieri''.


Description


''Camposipterus'' as a clade

Rodrigues & Kellner treated ''Camposipterus'' as a group or
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
. No
synapomorphies 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 have ...
could be established but a unique combination of themselves not unique traits was present. In side view the snout is rounded. The snout bears no crest. The front part of the snout is expanded. The palate curves upwards. The first tooth pair is located in the snout tip. Each of the species of ''Camposipterus'' has its own unique derived traits,
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 ...
, and a unique combination of traits.


''Camposipterus nasutus''

Rodrigues & Kellner established two autapomorphies of ''Camposipterus nasutus''. At the front of the jaw edge there is a density of three teeth per , at the rear a density of 2.5 teeth. The snout tip is flat, in front view wider than tall. There is a unique combination of traits: the upper profile of the snout is straight or lightly curved; the midline ridge on the palate is extended forwards until the level of the rear margins of the second tooth pair; to the rear the distance between the teeth gradually increases; the second and third tooth pairs are obliquely pointed sideways; the front of the snout is slightly expanded.


''Camposipterus''(?) ''sedgwickii''

Rodrigues & Kellner established two autapomorphies of ''Camposipterus''(?) ''sedgwickii''. The expanded section of the front snout is suddenly constricted behind the third tooth pair. The tooth sockets of the third tooth pair are much larger than those of the fourth pair. There is a unique combination of traits: the snout is deep; the midline ridge on the palate towards the front reaches a position behind the third tooth pair. This species was reassigned to the genus '' Aerodraco'' in 2020.


''Camposipterus''(?) ''colorhinus''

Rodrigues & Kellner established a single autapomorphy of ''Camposipterus''(?) ''colorhinus'': the depression above the first tooth pair is obliquely directed to below. There is a unique combination of traits: the frontal expansion of the snout is well-developed, without a constriction at its end; there is a depression above the first tooth pair; the second and third tooth sockets have a large diameter; the fourth tooth socket is much smaller than the second and third. ''Camposipterus''(?) ''colorhinus'' has a robust snout tip forming a large rosette to catch slippery prey such as fishes, indicating an animal of considerable size. The depression above the first tooth pair is shared with ''
Uktenadactylus ''Uktenadactylus'' is a genus of anhanguerid pterodactyloid pterosaurs from the Lower Cretaceous Paw Paw Formation of Texas, United States and the Wessex Formation on the Isle of Wight, England. Fossil remains of ''Uktenadactylus'' dated back to ...
wadleighi'', but in this form the depression is horizontally oriented. Seeley speculated that it was the attachment area for a lip but pterosaurs had horny beaks.


Phylogeny

Rodrigues and Kellner assigned ''Camposipterus'' to the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Anhangueria Anhangueria (or anhanguerians) is a group of pterosaurs belonging to the clade Pteranodontoidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stages), around 140 to 92.5 million years ...
, but ''
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, uncertainty ...
'', thus in an unknown position. Some
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analyses published in the naming article suggested a position in the evolutionary tree above ''Cimoliopterus'' and below ''
Cearadactylus ''Cearadactylus'' is a genus of large anhanguerid pterodactyloid pterosaur from the Romualdo Formation of Brazil, South America. Fossil remains of ''Cearadactylus'' dated back to the Albian stage of the Early Cretaceous period, about 112 million ...
atrox''. In 2019, Jacobs ''et al.'' published a
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
that placed ''Camposipterus'' within the family
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 ...
, more specifically the sister taxon of ''Cimoliopterus''. In the same year however, a study by Pêgas ''et al.'' placed ''Camposipterus'' within the clade
Targaryendraconia Targaryendraconia is an extinct clade of lanceodontian pterosaurs that lived from the Early to Late Cretaceous period in Europe, North America, South America, and Australia. Classification Below is a cladogram following a topology by Pêga ...
, and specifically within the family Cimoliopteridae as the sister taxon of both ''
Aetodactylus ''Aetodactylus'' (meaning "eagle finger") is a genus of targaryendraconian pterosaur. It is known from a lower jaw discovered in Upper Cretaceous rocks of northeastern Texas, United States. Description ''Aetodactylus'' is based on SMU 763 ...
'' and ''Cimoliopterus'':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, Topology 1: Jacobs ''et al.'' (2019). Topology 2: Pêgas ''et al.'' (2019).


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, United Kingdom Pteranodontoids Albian life Early Cretaceous pterosaurs of Europe Cretaceous England Fossils of England Fossil taxa described in 2013 Taxa named by Alexander Kellner