Thyreophora ("shield bearers", often known simply as "armored dinosaurs") is a group of
armored ornithischia
Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
n
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 that lived from the Early
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
until 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 ...
.
Thyreophorans are characterized by the presence of body armor lined up in longitudinal rows along the body. Primitive forms had simple, low, keeled scutes or
osteoderm
Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amph ...
s, whereas more derived forms developed more elaborate structures including spikes and plates. Most thyreophorans were
herbivorous
A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
and had relatively small brains for their body size.
Thyreophora includes two major subgroups,
Ankylosauria
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 limbs ...
and
Stegosauria
Stegosauria is a group of Herbivore, herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous Period (geology), periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere (North America, Europe a ...
. In both clades, the forelimbs were much shorter than the hindlimbs, particularly in stegosaurs. Thyreophora has been defined as the group consisting of all species more closely related to ''
Ankylosaurus'' and ''
Stegosaurus
''Stegosaurus'' (; ) is a genus of herbivorous, four-legged, armored dinosaur from the Late Jurassic, characterized by the distinctive kite-shaped upright plates along their backs and spikes on their tails. Fossils of the genus have been fo ...
'' than to ''
Iguanodon
''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species found worldwide have been classified in the genus ''Iguanodon'', dating from the Late Jurassic to Early Cretaceous, Taxonomy (bi ...
'' and ''
Triceratops
''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 ...
''. It is the
sister group
In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree.
Definition
The expression is most easily illustrated by a cladogram:
Taxon A and ...
of
Cerapoda
Neornithischia ("new ornithischians") is a clade of the dinosaur order Ornithischia. It is the sister group of the Thyreophora within the clade Genasauria. Neornithischians are united by having a thicker layer of asymmetrical enamel on the ins ...
within
Genasauria
Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek st ...
.
Characteristics
Members of Thyreophora are characterised by the presence of
osteoderms
Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of Extant taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...
(bony growths within the skin), with these osteoderms having lateral keels. Characters of the skull and jaws distinctive (
synapomorphic) of thyreophorans include "absence of a deep elliptic
fossa along the
sutural line of the
nasals, presence of a wide
jugal, remodeling of skull
dermal bone
A dermal bone or investing bone or membrane bone is a bony structure derived from intramembranous ossification forming components of the vertebrate skeleton, including much of the skull, jaws, gill covers, shoulder girdle, fin rays ( lepidotrich ...
, down-turned
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 ...
tooth row". Among primitive thyreophorans, ''
Scutellosaurus'' was likely primarily bipedal, while the more quadrupedally adapted ''
Scelidosaurus'' may have been bipedal for some of the time, particularly as a juvenile. Stegosaurs and ankylosaurs are thought to have been obligately quadrupedal.
Classification
Taxonomy
While ranked taxonomy has largely fallen out of favor among dinosaur paleontologists, a few 21st century publications have retained the use of ranks, though sources have differed on what its rank should be. Most have listed Thyreophora as an unranked taxon containing the traditional suborders Stegosauria and Ankylosauria, though Thyreophora is also sometimes classified as a suborder, with Ankylosauria and Stegosauria as infraorders.
Phylogeny
Thyreophora was first named by
Nopcsa in 1915. Thyreophora was defined as a
clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
by
Paul Sereno
Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago who has discovered several new dinosaur species on several continents, including at sites in Inner Mongolia, Argentina, Morocco and Niger. ...
in 1998, as "all
genasaurs more closely related to ''
Ankylosaurus'' than to ''
Triceratops
''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 ...
''". Thyreophoroidea was first named by Nopcsa in 1928 and defined by Sereno in 1986, as "''
Scelidosaurus'', ''
Ankylosaurus'', their most recent common ancestor and all of its descendants". Eurypoda was first named by Sereno in 1986 and defined by him in 1998, as "''
Stegosaurus
''Stegosaurus'' (; ) is a genus of herbivorous, four-legged, armored dinosaur from the Late Jurassic, characterized by the distinctive kite-shaped upright plates along their backs and spikes on their tails. Fossils of the genus have been fo ...
'', ''Ankylosaurus'', their most recent common ancestor and all of their descendants".
In 2021, an international group of researchers led by Daniel Madzia registered almost all of the most commonly used ornithischian clades under the
International Code of Phylogenetic Nomenclature, with the intent of standardizing their definitions. According to Madzia ''et al.'', Thyreophora is defined as the largest clade containing ''Ankylosaurus magniventris'' and ''Stegosaurus stenops'' but not ''
Iguanodon
''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species found worldwide have been classified in the genus ''Iguanodon'', dating from the Late Jurassic to Early Cretaceous, Taxonomy (bi ...
bernissartensis'' and ''
Triceratops
''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 ...
horridus''.
They also defined the less inclusive Eurypoda as "the smallest clade containing ''
Ankylosaurus magniventris'' and ''
Stegosaurus stenops''" to include the ankylosaurs and stegosaurs to the exclusion of basal thyreophorans. A later study conducted by André Fonseca and colleagues in 2024 gave a formal definition for Thyreophoroidea in 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 ...
'' as "the smallest clade containing ''
Ankylosaurus magniventris'', ''
Scelidosaurus harrisonii'', and ''
Stegosaurus stenops''".
The following
cladogram
A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
shows the results of the
phylogenetic analysis
In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
Soto-Acuña ''et al''. (2021).
In their description of ''
Jakapil'' the following year, Riguetti ''et al'' modified the same matrix and found it to occupy a position as the
sister taxon
In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree.
Definition
The expression is most easily illustrated by a cladogram:
Taxon A and ...
to the Eurypoda.
A similar result was found by Fonseca et al. in 2024.
In 2020, as part of his monograph on ''
Scelidosaurus'',
David Norman revised the relationships of early thyreophorans, finding that Stegosauria was the most basal branch, with ''
Scutellosaurus'', ''
Emausaurus'' and ''Scelidosaurus'' being progressive
stem groups to Ankylosauria, rather than to Stegosauria+Ankylosauria. A cladogram is given below:
See also
* ''
thyreophoroi'', Greek soldiers bearing a
thyreos shield
*
List of thyreophoran type specimens
References
{{Taxonbar, from=Q823931
Dinosaur clades