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''Europejara'' is a genus of
tapejarid Tapejaridae (from a Tupi word meaning "the old being") are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primiti ...
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 ...
period of
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
. The type and only species known is ''Europejara olcadesorum''.


Discovery and naming

In 2012, 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 ...
''Europejara olcadesorum'' was named and described by Romain Vullo, Jesús Marugán-Lobón,
Alexander Kellner Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...
, Angela Buscalioni, Bernard Gomez, Montserrat de la Fuente and José Moratalla. The generic name combines the names of
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and the related genus '' Tapejara'', in reference to the fact that ''Europejara'' is the first tapejarid found in that continent. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
refers to the
Olcades The Olcades were an ancient stock-raising pre-Roman people from Hispania, who lived to the west of the Turboletae in the southeastern fringe of the Iberian system mountains. Origins Related to both the Celtiberians and Carpetani, the Olcades ap ...
, the Celtiberic tribe inhabiting the region of Cuenca, the location of the find, in
Antiquity Antiquity or Antiquities may refer to: Historical objects or periods Artifacts *Antiquities, objects or artifacts surviving from ancient cultures Eras Any period before the European Middle Ages (5th to 15th centuries) but still within the histo ...
. 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 ...
, MCCM-LH 9413, was uncovered at the Las Hoyas site in a chalkstone layer of the Calizas de La Huergina Formation dating from the late
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 preceded ...
. It consists of a partial skull with lower jaws, compressed on a slab and counterslab. Two elements of the
hyoid The hyoid bone (lingual bone or tongue-bone) () is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid cartilage. At rest, it lies between the base of the mandible and the third cervical vertebra. ...
are present also. The skull has been vertically crushed, the lower jaws horizontally. The specimen was prepared by Mercedes Llandres Serrano, and is part of the Las Hoyas collection of the Museo de las Ciencias de Castilla–La Mancha.


Description

''Europejara'' is a relatively small form with an estimated
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 of ...
of . The jaws are toothless and the lower jaws bear a large downwards pointing crest. The describers established three
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 ...
, unique derived traits. The crest on the lower jaws is curved to the back. The crest is deeper than its base, measured from the front to the back, is wide. The crest is four times deeper than the back of the jaw. Two other diagnostic traits were indicated: the inner side of the lower jaw is thickened, causing a convex curvature; the inner side shows some shallow, but well-demarcated, depressions. Due to the crushing of the skull, its fragments, mainly representing elements from the area around the right eye socket, show little detail. The lower jaws have a preserved length of and an estimated original length of . In their front parts the lower jaws are fused by a symphysis into a
mandibula In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
. The symphysis has a concave upper profile and features a large crest on the underside, pointing downwards for at least . The back edge of the crest is recurved; the curvature of the front edge cannot be exactly established because of damage. The crest is the longest relative to lower jaw length of any known pterosaur. The internal bone structure of the crest is spongy. The rod-like first ceratobranchialia pair of the hyoid have a length of and a cross-section of .


Phylogeny

''Europejara'' was assigned to the
Tapejaridae Tapejaridae (from a Tupi word meaning "the old being") are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primiti ...
. A
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 ...
analysis showed it to be more precisely a member of the
Tapejarinae Tapejaridae (from a Tupi word meaning "the old being") are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primiti ...
. Apart from being the first tapejarid known from Europe, it would also be the oldest pterosaur with certainty known to be edentulous; older fragments have been reported representing other generally toothless
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, ...
s but these did not include the jaws themselves. The cladogram below follows 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 ...
by Kellner, one of the describers of ''Europejara'', and colleagues in 2019. They recovered ''Europejara'' within the tribe
Tapejarini Tapejaridae (from a Tupi word meaning "the old being") are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primiti ...
, sister taxon to three other genera: ''
Caiuajara ''Caiuajara'' is an extinct genus of tapejarid pterosaur from the Late Cretaceous period ( Santonian stage) of Brazil. It is known from a single type species, ''Caiuajara dobruskii''. Discovery and naming In 1971, the labourers Alexandre Dobr ...
'', '' Tapejara'', and ''
Tupandactylus ''Tupandactylus'' (meaning "Tupan finger", in reference to the Tupi thunder god) is a genus of tapejarid pterodactyloid pterosaur from the Early Cretaceous Crato Formation of Brazil. History ''Tupandactylus imperator'' is known from four nearl ...
''.


Paleobiology

Following earlier suggestions about the diet of tapejarids, the describers assumed a
frugivorous A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance and ...
lifestyle for ''Europejara''. Because the species is so old it indicates a role for the tapejarids in the
Cretaceous Terrestrial Revolution The Cretaceous Terrestrial Revolution (abbreviated KTR), also known as the Angiosperm Terrestrial Revolution (ATR) by authors who consider it to have lasted into the Palaeogene, describes the intense diversification of angiosperms, insects, reptiles ...
, a turn-over in the
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s of the Lower Cretaceous in which
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, Î³Ï…Î¼Î½ÏŒÏ ...
s were replaced by
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
, flowering plants, and new groups of herbivores evolved, adapted to the changed food supply. In the case of tapejarids there could have been a reinforcing interactive cycle between the evolution of fruit and the pterosaurs dispersing the seed. Possibly the beaks of the tapejarids had ragged edges forming pseudo-teeth to better separate the fruit flesh from the seeds, as with some extant
toucans Toucans (, ) are members of the Neotropical near passerine bird family Ramphastidae. The Ramphastidae are most closely related to the American barbets. They are brightly marked and have large, often colorful bills. The family includes five gen ...
.


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, Spain Tapejaromorphs Early Cretaceous pterosaurs of Europe Cretaceous Spain Fossils of Spain La Huérguina Formation Fossil taxa described in 2012 Taxa named by Alexander Kellner