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Leptoceratopsidae is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
of neoceratopsian
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s from
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
,
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
and
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 ...
. Leptoceratopsids resembled, and were closely related to, other neoceratopsians, such as the families Protoceratopsidae and Ceratopsidae, but they were more primitive and generally smaller.


Phylogeny

Leptoceratopsidae was originally named by Franz Nopcsa von Felső-Szilvás in 1923 as a subfamily Leptoceratopsinae, and its type species is '' Leptoceratops gracilis''. Mackovicky, in 2001, defined it as a stem-based taxon and a family consisting of ''Leptoceratops gracilis'' and all species closer to ''Leptoceratops'' than to '' Triceratops horridus''.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. The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
below follows the topologies from a 2015 analysis by Yiming He, Peter J. Makovicky, Kebai Wang, Shuqing Chen, Corwin Sullivan, Fenglu Han, Xing XuMichael J. Ryan, David C. Evans, Philip J. Currie, Caleb M. Brown and Don Brinkman.


Palaeobiogeography

Definitive leptoceratopsids have so far been found exclusively in the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger 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 ''creta'', ...
period (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 mya. ...
- late
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the inte ...
stages) of
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
and Western
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
; 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. Campani ...
of
North Carolina North Carolina () is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. The state is the List of U.S. states and territories by area, 28th largest and List of states and territories of the United ...
and
Sweden Sweden, formally the Kingdom of Sweden,The United Nations Group of Experts on Geographical Names states that the country's formal name is the Kingdom of SwedenUNGEGN World Geographical Names, Sweden./ref> is a Nordic countries, Nordic c ...
extends their geographic range into Eastern North America and
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 ...
. A possible leptoceratopsid
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
, named ''
Serendipaceratops ''Serendipaceratops'' (meaning "serendipitous horned face") is a genus of herbivorous ornithischian dinosaur, possibly an ankylosaur, from the early Cretaceous Period of Australia. The type species, ''S. arthurcclarkei'', was named in 2003. Di ...
'', has been found in Victoria, Australia. However, a 2010 study showed that it could not be confidently referred to any ornithischian family, and 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 s ...
''.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 mya. ...
, to ''
Leptoceratops ''Leptoceratops'' (meaning 'Thin-horned face' and derived from Greek ''lepto-/λεπτο-'' meaning 'small', 'insignificant', 'slender', 'meagre' or 'lean', ''kerat-/κερατ-'' meaning 'horn' and ''-ops/ωψ'' meaning face), is a genus of l ...
'', right at the end of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
in the late
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the inte ...
. ''Gryphoceratops'' is the first definitive record of Santonian leptoceratopsid. It was named based on a partial left dentary from
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest T ...
,
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by to ...
. ''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 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 ...
dated to the early Campanian of North Carolina, whilst the European material referred to Leptoceratopsidae consists of isolated teeth and
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...
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 cultural region in the Eastern United States that stretches from the Southern Tier of New York State to northern Alabama and Georgia. While the Appalachian Mountains stretch from Belle Isle in Newfoundland and Labrador, C ...
was isolated from Western Europe and Laramidia 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 The Baltic Shield (or Fennoscandian Shield) is a segment of the Earth's crust belonging to the East European Craton, representing a large part of Fennoscandia, northwestern Russia and the northern Baltic Sea. It is composed mostly of Archean an ...
was also an isolated landmass during the Late Cretaceous.


See also

* Timeline of ceratopsian research


References

{{Taxonbar, from=Q123245 Cretaceous dinosaurs Dinosaurs of Asia Dinosaurs of North America Taxa named by Franz Nopcsa von Felső-Szilvás Prehistoric dinosaur families