Leptoceratopsidae is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

neoceratopsian Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period, although ancestral forms lived earlier, in the Late Ju ...

dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...

s from

Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

and possibly

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

. Leptoceratopsids resembled, and were closely related to, other neoceratopsians, such as the

families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...

Protoceratopsidae Protoceratopsidae is a family of basal (primitive) ceratopsians from the Late Cretaceous period. Although ceratopsians have been found all over the world, protoceratopsids are only definitively known from Cretaceous strata in Asia, with most spec ...

and

Ceratopsidae Ceratopsidae (sometimes spelled Ceratopidae) is a family of ceratopsian dinosaurs including ''Triceratops'', ''Centrosaurus'', and ''Styracosaurus''. All known species were quadrupedal herbivores from the Upper Cretaceous. All but one species are k ...

, but they were more primitive and generally smaller.

Phylogeny

Leptoceratopsidae was originally named by

Franz Nopcsa von Felső-Szilvás Baron Franz Nopcsa von Felső-Szilvás (also Baron Nopcsa von Felső-Szilvás, Baron Nopcsa, Ferenc Nopcsa, báró felsőszilvási Nopcsa Ferenc, Baron Franz Nopcsa, and Franz Baron Nopcsa; May 3, 1877 – April 25, 1933) was a Hungarian Aristoc ...

in 1923 as a subfamily Leptoceratopsinae, and its type species is ''

Leptoceratops gracilis ''Leptoceratops'' (meaning 'small horn face') is a genus of ceratopsian dinosaur from the Late Cretaceous of North America. First found in Alberta in 1910, the type species ''Leptoceratops gracilis'' was named in 1914 by Barnum Brown for a parti ...

''. Mackovicky, in 2001, defined it as a

stem-based taxon Phylogenetic nomenclature is a method of nomenclature for taxon, taxa in biology that uses phylogenetics, phylogenetic definitions for taxon names as explained below. This contrasts with Biological classification, the traditional method, by which ...

and a family consisting of ''Leptoceratops gracilis'' and all species closer to ''Leptoceratops'' than to ''

Triceratops horridus ''Triceratops'' ( ; ) is a genus of Chasmosaurinae, chasmosaurine Ceratopsia, ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous Period (geology), period, about 68 to 66 million years ago on the island ...

''.Makovicky, P.J. 2001. A ''Montanoceratops cerorhynchus'' (Dinosauria: Ceratopsia) braincase from the Horseshoe Canyon Formation of Alberta, In: Tanke, D.H. & Carpenter, K. (Eds.). ''Mesozoic Vertebrate Life''. Bloomington: Indiana University Press. Pp. 243-262. All previously published neoceratopsian phylogenetic analyses were incorporated into the analysis of Eric M. Morschhauser and colleagues in

2019 This was the year in which the first known human case of COVID-19 was documented, preceding COVID-19 pandemic, the pandemic which was declared by the World Health Organization the following year. Up to that point, 2019 had been described as ...

, along with all previously published diagnostic species excluding the incomplete juvenile '' Archaeoceratops yujingziensis'' and the problematic genera '' Bainoceratops'', '' Lamaceratops'', '' Platyceratops'' and '' Gobiceratops'' that are very closely related to and potentially synonymous with ''

Bagaceratops ''Bagaceratops'' (meaning "small-horned face") is a genus of small protoceratopsid dinosaurs that lived in Asia during the Late Cretaceous, around 72 to 71 million years ago. ''Bagaceratops'' remains have been reported from the Barun Goyot Forma ...

''. While there were many unresolved areas of the strict consensus, including all of Leptoceratopsidae, a single most parsimonious tree was found that was most consistent with the relative ages of the taxa included, which is shown below.

Palaeobiogeography

Definitive leptoceratopsids lived in

and Western

during the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...

around 83-66 million years ago. However, material referred to leptoceratopsids from the early

Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...

North Carolina North Carolina ( ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It is bordered by Virginia to the north, the Atlantic Ocean to the east, South Carolina to the south, Georgia (U.S. stat ...

and

Sweden Sweden, formally the Kingdom of Sweden, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. It borders Norway to the west and north, and Finland to the east. At , Sweden is the largest Nordic count ...

possibly extends their geographic range into Eastern North America and

. A possible leptoceratopsid

ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...

named '' Serendipaceratops'' has been found in Australia, but this taxon is considered a ''

nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...

'' due to the fragmentary holotype which cannot be confidently referred to any ornithischian group, though it might possibly belong to an

ankylosaur Ankylosauria is a group of herbivorous dinosaurs of the clade Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful l ...

instead.Agnolin, F.L., Ezcurra, M.D., Pais, D.F. and Salisbury, S.W. (2010). "A reappraisal of the Cretaceous non-avian dinosaur faunas from Australia and New Zealand: Evidence for their Gondwanan affinities." ''Journal of Systematic Palaeontology'', 8(2): 257-300. Leptoceratopsids range in age from '' Gryphoceratops'', of the late

Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya ( million years ago) and 83.6 ± 0.7 m ...

, to ''

Leptoceratops ''Leptoceratops'' (meaning 'small horn face') is a genus of ceratopsian dinosaur from the Late Cretaceous of North America. First found in Alberta in 1910, the type species ''Leptoceratops gracilis'' was named in 1914 by Barnum Brown for a parti ...

'', right at the end of the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

in the late

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

. ''Gryphoceratops'' is the first definitive record of Santonian leptoceratopsid. It was named based on a partial left

dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...

from

Alberta Alberta is a Provinces and territories of Canada, province in Canada. It is a part of Western Canada and is one of the three Canadian Prairies, prairie provinces. Alberta is bordered by British Columbia to its west, Saskatchewan to its east, t ...

Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...

. ''Gryphoceratops'' represents the oldest known leptoceratopsid and probably the smallest adult ceratopsian known from North America. Leptoceratopsids are known from Eastern North America by a partial

maxilla In vertebrates, the maxilla (: maxillae ) 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. The two maxil ...

dated to the early Campanian of North Carolina, whilst the European material referred to Leptoceratopsidae consists of isolated teeth and

vertebrae Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...

from the early Campanian of Sweden. The former represents the first known ceratopsian from Eastern North America, and its specialised maxillary anatomy supports the hypothesis that

Appalachia Appalachia ( ) is a geographic region located in the Appalachian Mountains#Regions, central and southern sections of the Appalachian Mountains in the east of North America. In the north, its boundaries stretch from the western Catskill Mountai ...

was isolated from Western Europe and

Laramidia Laramidia was an island continent that existed during the Late Cretaceous period (99.6–66 Year#SI prefix multipliers, Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island ...

for an extended period during the Late Cretaceous, resulting in an endemic Late Cretaceous fauna. The shared presence of leptoceratopsids in Appalachia and Western Europe has implications for their biogeographic dispersal, suggesting the possibility that leptoceratopsids entered Appalachia through either Western Europe or Laramidia; however, it is also possible that the European leptoceratopsids also represented a distinct endemic assemblage, as the Fennoscandian Shield was also an isolated landmass during the Late Cretaceous.

References

{{Taxonbar, from=Q123245 Late Cretaceous dinosaurs Dinosaur families

Phylogeny

Palaeobiogeography

See also

References