Polacanthidae
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Polacanthinae is a subfamily of
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 ...
s, most often
nodosaurids Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous Period (geology), periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophylet ...
, from the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
through
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
of
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 ...
and potentially
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
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 ...
. The group is defined as the largest clade closer to ''
Polacanthus foxii ''Polacanthus'', deriving its name from the Ancient Greek polys-/πολύς- "many" and akantha/ἄκανθα "thorn" or "prickle", is an early armoured, spiked, plant-eating ankylosaurian dinosaur from the early Cretaceous period of England. In ...
'' than ''
Nodosaurus textilis ''Nodosaurus'' (meaning 'knobbed lizard') is a genus of herbivore, herbivorous nodosauridae, nodosaurid ankylosaurian dinosaur from the Late Cretaceous, the fossils of which are found exclusively in the Frontier Formation in Wyoming. Descripti ...
'' or ''
Ankylosaurus magniventris ''Ankylosaurus'' is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of th ...
'', as long as that group nests within either Nodosauridae or
Ankylosauridae Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
. If ''Polacanthus'', and by extent Polacanthinae, falls outside either family-level clade, then the ''-inae'' suffix would be inappropriate, and the proper name for the group would be the informally defined Polacanthidae. Polacanthines were somewhat more lightly armoured than more advanced ankylosaurids and nodosaurids. Their spikes were made up of thin, compact bone with less reinforcing collagen than in the heavily armoured nodosaurids. The relative fragility of polacanthine armour suggests that it may have been as much for display as defense. They appear to have become extinct about the same time a
land bridge In biogeography, a land bridge is an isthmus or wider land connection between otherwise separate areas, over which animals and plants are able to cross and colonize new lands. A land bridge can be created by marine regression, in which sea le ...
opened between Asia and North America.Kirkland, J. I. (1996). Biogeography of western North America's mid-Cretaceous faunas - losing European ties and the first great Asian-North American interchange. J. Vertebr. Paleontol. 16 (Suppl. to 3): 45A.


Classification

The family Polacanthidae was named by Ernst Jaekel in
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to refer to a group of ankylosaurs which seemed to him intermediate between the ankylosaurids and nodosaurids. This grouping was ignored by most researchers until the late 1990s, when it was used as a subfamily (Polacanthinae) by Kirkland for a natural group recovered by his 1998 analysis suggesting that ''
Polacanthus ''Polacanthus'', deriving its name from the Ancient Greek polys-/πολύς- "many" and akantha/ἄκανθα "thorn" or "prickle", is an early armoured, spiked, plant-eating ankylosaurian dinosaur from the early Cretaceous period of England. In ...
'', '' Gastonia'', and ''
Mymoorapelta ''Mymoorapelta'' (Meaning "Shield from the Mygatt-Moore Quarry" after a combination of the names of the discoverers of the Mygatt-Moore Quarry that fossils were originally collected from, and the Greek word πέλτα, meaning "shield") is a Nod ...
'' were closely related within the family Ankylosauridae, and he additionally referred ''
Hoplitosaurus ''Hoplitosaurus'' (meaning "Hoplite lizard") was a genus of armored dinosaur related to ''Polacanthus''. It was named from a partial skeleton found in the Barremian-age Lower Cretaceous Lakota Formation of Custer County, South Dakota. It is an ...
'' and ''
Hylaeosaurus ''Hylaeosaurus'' ( ; Greek: / "belonging to the forest" and / "lizard") is a herbivorous ankylosaurian dinosaur that lived about 136 million years ago, in the late Valanginian stage of the early Cretaceous period of England. It was found i ...
'' to the subfamily, though they were too incomplete to analyze. Kenneth Carpenter resurrected the name Polacanthidae for a similar group which he also found to be closer to ankylosaurids than to nodosaurids. Carpenter became the first to define Polacanthidae as all dinosaurs closer to ''Gastonia'' than to either ''
Edmontonia ''Edmontonia'' is a genus of panoplosaurin nodosaurid dinosaur from the Late Cretaceous Period. It is part of the Nodosauridae, a family within Ankylosauria. It is named after the Edmonton Formation (now the Horseshoe Canyon Formation in Canada ...
'' or ''
Euoplocephalus ''Euoplocephalus'' ( ) is a genus of large herbivorous ankylosaurid dinosaurs, living during the Late Cretaceous of Canada. It has only one named species, ''Euoplocephalus tutus''. The first fossil of ''Euoplocephalus'' was found in 1897 in Albe ...
''. Most subsequent researchers placed polacanthines as primitive ankylosaurids, though mostly without any rigorous study to demonstrate this idea. The first comprehensive study of 'polacanthid' relationships, published in 2011, found that they are either an unnatural grouping of primitive nodosaurids, or a valid subfamily at the base of Nodosauridae. In the act of formally defining Polacanthinae in line with the requirements of the
PhyloCode The ''International Code of Phylogenetic Nomenclature'', known as the ''PhyloCode'' for short, is a formal set of rules governing phylogenetic nomenclature. Its current version is specifically designed to regulate the naming of clades, leaving the ...
, Madzia and colleagues chose for the analysis of Yang ''et al.'' (2013) to be the primary reference for Polacanthinae, supplemented by similar results found by Kirkland (1998), Thompson ''et al.'' (2012), Arbour ''et al.'' (2016), Rivera-Sylva ''et al.'' (2018), and Zheng ''et al.'' (2018). While the clade is frequently referenced in literature, a majority of taxa considered to be polacanthines have instead been found to not form any natural groups with ''Polacanthus'', representing both primitive nodosaurids, primitive ankylosaurids, or outside the Nodosauridae-Ankylosauridae clade. In the analysis of Arbour ''et al.'' in
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, and its subsequent modifications by Rivera-Sylva ''et al.'' and Zheng ''et al.'' in
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, only ''Hoplitosaurus'' was resolved as a polacanthine alongside ''Polacanthus'', with ''Peloroplites'' and ''Taohelong'' falling within Nodosaurinae, ''Gargoyleosaurus'', ''Gastonia'', and ''Mymoorapelta'' being more basal nodosaurids outside both subclades, and ''Hylaeosaurus'' representing an early-diverging ankylosaurid.


References

{{Taxonbar, from=Q21711005 Dinosaur subfamilies Early Cretaceous dinosaurs