Ankylosauria is a group of
herbivorous
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
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 of the order
Ornithischia
Ornithischia () is an extinct order 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 s ...
. It includes the great majority of dinosaurs with
armor
Armour (British English) or armor (American English; see spelling differences) is a covering used to protect an object, individual, or vehicle from physical injury or damage, especially direct contact weapons or projectiles during combat, or fr ...
in the form of bony osteoderms, similar to
turtles
Turtles are an order (biology), order of reptiles known as Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) an ...
. Ankylosaurs were bulky
quadruped
Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
s, with short, powerful limbs. They are known to have first appeared in the
Middle Jurassic
The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
, and persisted until the end of the
Cretaceous Period
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 t ...
. The two main families of Ankylosaurs,
Nodosauridae
Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia.
Description
Nodosaurids, like their close relatives the ankylosaurids, wer ...
and
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 ...
are primarily known from the Northern Hemisphere, but the more basal
Parankylosauria
Parankylosauria is a group of basal ankylosaurian dinosaurs known from the Cretaceous of South America, Antarctica, and Australia. It is thought the group split from other ankylosaurs during the mid-Jurassic period, despite this being unpreserved ...
are known from southern
Gondwana
Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
during the Cretaceous.
Ankylosauria was first named by
Henry Fairfield Osborn
Henry Fairfield Osborn, Sr. (August 8, 1857 – November 6, 1935) was an American paleontologist, geologist and eugenics advocate. He was the president of the American Museum of Natural History for 25 years and a cofounder of the American Euge ...
in 1923.
[Osborn, H. F. (1923). "Two Lower Cretaceous dinosaurs of Mongolia." ''American Museum Novitates'', 95: 1–1]
/ref> In the Linnaean classification
Linnaean taxonomy can mean either of two related concepts:
# The particular form of Taxonomy (biology), biological classification (taxonomy) set up by Carl Linnaeus, as set forth in his ''Systema Naturae'' (1735) and subsequent works. In the ta ...
system, the group is usually considered either a suborder or an infraorder. It is contained within the group Thyreophora
Thyreophora ("shield bearers", often known simply as "armored dinosaurs") is a group of armored ornithischian dinosaurs that lived from the Early Jurassic until the end of the Cretaceous.
Thyreophorans are characterized by the presence of body ...
, which also includes the stegosaur
Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe, ...
s, armored dinosaurs known for their combination of plates and spikes.
Etymology
The name of this group of dinosaurs is associated with a number of anatomical features in which small and large bony shields fused together, completely covering their back and sides. On the skull these shields fused with the underlying bones, and the dorsal ribs fit snugly to the vertebrae. The Latin name Ankylosauria is derived from the Greek ἀγκύλος nkylos— "curved", "bent" with the anatomical meaning "hard" or "fused" and σαῦρος auros
Auros (; oc-gsc, Auròs) is a commune in the Gironde department in southwestern France.
Population
See also
*Communes of the Gironde department
The following is a list of the 535 communes of the Gironde department of France.
The commune ...
— "lizard".[Ben Creisler, ''Dinosauria Translation and Pronunciation Guide.'']
In the 1908 description of the genus Ankylosaurus
''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 the n ...
, Barnum Brown
Barnum Brown (February 12, 1873 – February 5, 1963), commonly referred to as Mr. Bones, was an American paleontologist. Named after the circus showman P. T. Barnum, he discovered the first documented remains of ''Tyrannosaurus'' during a career ...
described the family 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 ...
as a group of representatives with a "rigid spine", but noted the wide, curved shape of the ribs, suggesting a "strongly curved" back (an error based on the alleged similarity to stegosaurs
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, predominantly in what i ...
and glyptodonts
Glyptodonts are an extinct subfamily of large, heavily armoured armadillos. They arose in South America around 48 million years ago and spread to southern North America after the continents became connected several million years ago. The best-k ...
, as ankylosaurs have flat backs). Therefore, "rigid lizard" and "curved lizard" could be additional meanings applied to the name of ankylosaurs.[
]
Classification
Ankylosauria and Stegosauria
Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe, ...
together form the two major subgroups of Thyreophora
Thyreophora ("shield bearers", often known simply as "armored dinosaurs") is a group of armored ornithischian dinosaurs that lived from the Early Jurassic until the end of the Cretaceous.
Thyreophorans are characterized by the presence of body ...
, a group of armoured dinosaurs distinct from ornithopod
Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous wo ...
s and marginocephalia
Marginocephalia (/mär′jə-nō-sə-făl′ē-ən/ Latin: margin-head) is a clade of ornithischian dinosaurs that is characterized by a bony shelf or margin at the back of the skull. These fringes were likely used for display. There are two clad ...
ns. Historically used for forms lacking large vertical plates, Kenneth Carpenter
Kenneth Carpenter (born September 21, 1949, in Tokyo, Japan) is a paleontologist. He is the former director of the USU Eastern Prehistoric Museum and author or co-author of books on dinosaurs and Mesozoic life. His main research interests ...
proposed in 1997
File:1997 Events Collage.png, From left, clockwise: The movie set of ''Titanic'', the highest-grossing movie in history at the time; ''Harry Potter and the Philosopher's Stone'', is published; Comet Hale-Bopp passes by Earth and becomes one of t ...
the first informal definition of the group, as all ornithischians closer to ''Ankylosaurus
''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 the n ...
'' than ''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 foun ...
''.[Carpenter, K., 1997, "Ankylosauria" pp. 16-20 in: P.J. Currie and K. Padian (eds.), ''Encyclopedia of Dinosaurs'', Academic Press, San Diego] This definition was further refined by Paul Sereno
Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at sites ...
in 2005
File:2005 Events Collage V2.png, From top left, clockwise: Hurricane Katrina in the Gulf of Mexico; the Funeral of Pope John Paul II is held in Vatican City; "Me at the zoo", the first video ever to be uploaded to YouTube; Eris was discovered in ...
to specify ''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 the n ...
'' and ''Stegosaurus stenops
''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 foun ...
'', the type species of both genera, a definition that was followed by Madzia and colleagues in 2021
File:2021 collage V2.png, From top left, clockwise: the James Webb Space Telescope was launched in 2021; Protesters in Yangon, Myanmar following the 2021 Myanmar coup d'état, coup d'état; A civil demonstration against the October–November 2021 ...
when the group name and definition was formalized following 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 ...
. Phylogenetic and morphological studies have differed on the inclusion of certain early taxa into Ankylosauria, especially the armoured Early Jurassic
The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
form ''Scelidosaurus
''Scelidosaurus'' (; with the intended meaning of "limb lizard", from Greek / meaning 'rib of beef' and ''sauros''/ meaning 'lizard')Liddell & Scott (1980). Greek-English Lexicon, Abridged Edition. Oxford University Press, Oxford, UK. is a gen ...
''. As some analyses, like that of Carpenter from 2001
The September 11 attacks against the United States by Al-Qaeda, which Casualties of the September 11 attacks, killed 2,977 people and instigated the global war on terror, were a defining event of 2001. The United States led a Participants in ...
or David B. Norman in 2021 find ''Scelidosaurus'' and possibly other early forms like ''Emausaurus
''Emausaurus'' is a genus of thyreophoran or armored dinosaur from the Early Jurassic (Early Toarcian). Its fossils have been found in Mecklenburg-Vorpommern, northern Germany. ''Emausaurus'' is the only known Toarcian thyreophoran, as well as th ...
'' and ''Scutellosaurus
''Scutellosaurus'' ( ) is a genus of thyreophoran ornithischian dinosaur that lived approximately 196 million years ago during the early part of the Jurassic Period in what is now Arizona, USA. It is classified in Thyreophora, the armoured di ...
'' to fall closer to ''Ankylosaurus'' than ''Stegosaurus'', Carpenter and later Norman suggested redefining Ankylosauria to limit it to the two subclades Nodosauridae
Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia.
Description
Nodosaurids, like their close relatives the ankylosaurids, wer ...
and 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 ...
, creating the new clade Ankylosauromorpha for all taxa closer to ''Ankylosaurus'' than ''Stegosaurus''. However, as historically even these primitive forms were considered ankylosaurs if they were more derived than ''Stegosaurus'', Madzia and colleagues considered a redefinition of Ankylosauria to be undesirable, instead preferring to abandon Ankylosauromorpha as a same-definition junior synonym of Ankylosauria. The informal clade Euankylosauria, suggested by Norman as an alternative to redefining Ankylosauria, was discussed as potentially being useful in future for the typical group of nodosaurids and ankylosaurids, and was defined in 2021 by Soto-Acuña and colleagues as such as the result of their novel phylogenetic results, which grouped ''Antarctopelta
''Antarctopelta'' ( ; meaning 'Antarctic shield') was a genus of ankylosaurian dinosaur with one known species, ''A. oliveroi'', which lived in Antarctica during the Late Cretaceous Period. It was a medium-sized ankylosaur, reaching 4 meters (13& ...
'', ''Kunbarrasaurus
''Kunbarrasaurus'' is an extinct genus of small herbivorous ankylosaurian dinosaur from the Cretaceous of Australia.
Discovery
In November 1989, at Marathon Station near Richmond, Queensland, the skeleton was discovered of an ankylosaurian. ...
'' (''Minmi
Michiko Evwana (born December 8, 1974), better known by her stage name , is a Japanese hip-hop and reggae musician,, United Daily News (in Chinese), August 22, 2008 as well as a singer-songwriter and record producer. She is the first soca ar ...
'' sp. of earlier analyses) and their new taxon ''Stegouros
''Stegouros'' (, meaning "roofed tail") is a genus of ankylosaurian dinosaur from the Late Cretaceous Dorotea Formation of southern Chile. The genus contains a single species, ''Stegouros elengassen'', known from a semi-articulated, near-comple ...
'' as a unique group of non-euankylosaurian ankylosaurs, which they gave the name Parankylosauria
Parankylosauria is a group of basal ankylosaurian dinosaurs known from the Cretaceous of South America, Antarctica, and Australia. It is thought the group split from other ankylosaurs during the mid-Jurassic period, despite this being unpreserved ...
. The result of their phylogenetic analysis is below.
Evolution
The origin of ankylosaurs is poorly understood, and only a few specimens from the Middle Jurassic are known. The ancestry of ankylosaurs has long been sought among stegosaurs, the closest group to ankylosaurs compared to other dinosaurs. Currently, ankylosaurs are a close group of stegosaurs within the Eurypoda clade. They are united by the presence of osteoderms in the skin, the narrow triangular skull of stegosaurs is similar to that of nodosaurids, and some similarity is found in the structure of the palate. Since stegosaurs are known from the Middle Jurassic, ankylosaurs are probably of the same age. They may have split up during the Aalenian period, more than 170 million years ago, but they were definitely in Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
by the Bathonian
In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age.
Strat ...
due to the presence of ''Spicomellus
''Spicomellus'' (meaning "collar of spikes") is an extinct genus of herbivorous ankylosaurian dinosaur that lived in the supercontinent Gondwana during the Middle Jurassic Period. The type and only known species is ''Spicomellus afer'', named ...
'' in Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
. There are no well-preserved remains of ankylosaurs of that age. An incomplete radius and ulna from the Isle of Skye in Scotland are known, the exact affiliation of which to ankylosaurs or stegosaurs is not established. Most likely, ankylosaurs followed a different evolutionary path than stegosaurs, although it is unknown when and how they split off. In the latter, the osteoderms become raised, and the lateral protection disappears. Ankylosaurs evolved towards the development of osteoderms on the surface of the skull, increased armor and further consolidation of the carapace, which suggests that the ancestor of the carapace consisted of separate non-fused osteoderms.
Paleobiology
Possible neonate
An infant or baby is the very young offspring of human beings. ''Infant'' (from the Latin word ''infans'', meaning 'unable to speak' or 'speechless') is a formal or specialised synonym for the common term ''baby''. The terms may also be used to ...
-sized ankylosaur fossils have been documented in the scientific literature
: ''For a broader class of literature, see Academic publishing.''
Scientific literature comprises scholarly publications that report original empirical and theoretical work in the natural and social sciences. Within an academic field, scient ...
.[Tanke, D.H. and Brett-Surman, M.K. 2001. Evidence of Hatchling and Nestling-Size Hadrosaurs (Reptilia:Ornithischia) from Dinosaur Provincial Park (Dinosaur Park Formation: Campanian), Alberta, Canada. pp. 206-218. In: Mesozoic Vertebrate Life—New Research Inspired by the Paleontology of Philip J. Currie. Edited by D.H. Tanke and K. Carpenter. Indiana University Press: Bloomington. xviii + 577 pp.]
Armor
All ankylosaurians had armor over much of their bodies, mostly scutes and nodules, with large spines in some cases. The scutes, or plates, are rectangular to oval objects organized in transverse (side to side) rows, often with keels on the upper surface. Smaller nodules and plates filled in the open spaces between large plates. In all three groups, the first two rows of plates tend to form a sort of half-ring around the neck; in nodosaurids, this comes from adjacent plates fusing with each other (and there is a third row as well), while ankylosaurids usually have the plates fused to the top of another band of bone. The skull has armor plastered on to it, including a distinctive piece on the outside-rear of the lower jaw.
Diet and feeding
Ankylosaurs were built low to the ground, typically one foot off the ground surface. They had small, triangular teeth that were loosely packed, similar to stegosaurs
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, predominantly in what i ...
. The large hyoid bones left in skeletons indicates that they had long, flexible tongues. They also had a large, side secondary palate. This means that they could breathe while chewing, unlike crocodiles. Their expanded gut region suggests the use of fermentation to digest their food, using symbiotic bacteria and gut flora. Their diet likely consisted of fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s, cycads
Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male or ...
, and 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 ...
. Mallon et al. (2013) examined herbivore coexistence on the island continent of Laramidia
Laramidia was an island continent that existed during the Late Cretaceous period (99.6–66 Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island land mass separated from A ...
during the Late Cretaceous. It was concluded that ankylosaurs were generally restricted to feeding on vegetation at, or below, the height of 1 meter.
References
{{Taxonbar, from=Q273617
Bathonian first appearances
Maastrichtian extinctions
Taxa named by Henry Fairfield Osborn
Fossil taxa described in 1923