Anodontosaurus
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''Anodontosaurus'' 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 ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
ankylosaurid 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 ...
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 within the subfamily
Ankylosaurinae Ankylosaurinae is a subfamily of ankylosaurid dinosaurs, existing from the Early Cretaceous about 105 million years ago until the end of the Late Cretaceous, about 66 mya. Many genera are included in the clade, such as ''Ankylosaurus'', ''Pinacos ...
. It is known from the entire span of 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'', the ...
Horseshoe Canyon Formation The Horseshoe Canyon Formation is a stratigraphic unit of the Western Canada Sedimentary Basin in southwestern Alberta. It takes its name from Horseshoe Canyon, an area of badlands near Drumheller. The Horseshoe Canyon Formation is part of the ...
(mid Late
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. Campanian s ...
to "middle"
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 interval from ...
stage, about 72.8-67 Ma ago) of southern
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 Ter ...
,
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 tot ...
. It contains two
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, ''A. lambei'' and ''A. inceptus''.


Discovery

''Anodontosaurus'' was named by
Charles Mortram Sternberg Charles Mortram Sternberg (1885–1981) was an American-Canadian fossil collector and paleontologist, son of Charles Hazelius Sternberg. Late in his career, he collected and described ''Pachyrhinosaurus'', ''Brachylophosaurus'', ''Parksosaurus ...
in
1928 Events January * January – British bacteriologist Frederick Griffith reports the results of Griffith's experiment, indirectly proving the existence of DNA. * January 1 – Eastern Bloc emigration and defection: Boris Bazhanov, J ...
, based on
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
CMN 8530, a partially preserved
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
including the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
,
half ring A ring diacritic may appear above or below letters. It may be combined with some letters of the extended Latin alphabets in various contexts. Rings Distinct letter The character Å (å) is derived from an A with a ring. It is a distinct let ...
,
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 ...
and other
postcranial Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
remains.C. M. Sternberg (1929) "A toothless armoured dinosaur from the Upper Cretaceous of Alberta." ''Canada Department of Mines Geological Survey Bulletin (Geological Series)'' 54(49):28-33 The badly crushed skeleton was collected by Sternberg in 1916 from a
Canadian Museum of Nature The Canadian Museum of Nature (french: Musée canadien de la nature; CMN) is a national natural history museum based in Canada's National Capital Region. The museum's exhibitions and public programs are housed in the Victoria Memorial Museum Bui ...
quarry, 8 miles southwest of Morrin. It was collected from the upper part of the Lower
Horseshoe Canyon Formation The Horseshoe Canyon Formation is a stratigraphic unit of the Western Canada Sedimentary Basin in southwestern Alberta. It takes its name from Horseshoe Canyon, an area of badlands near Drumheller. The Horseshoe Canyon Formation is part of the ...
(unit 2), dating to the latest
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. Campanian s ...
to the earliest
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 interval from ...
stage of 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'', the ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
, about 71-70.2 
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
. The generic name means "toothless lizard" in
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
. It was inspired by the fact that compression damage to the specimen had removed the teeth, at the same time shifting various flat round elements below the skull and on top of the left lower jaw, misleading Sternberg into assuming that large "trituration plates" had replaced the normal dentition.Vickaryous, M.K. and Russell, A.P., 2003, "A redescription of the skull of ''Euoplocephalus tutus'' (Archosauria: Ornithischia): a foundation for comparative and systematic studies of ankylosaurian dinosaurs", ''Zoological Journal of the Linnean Society'' 137: 157–186 The specific name, ''lambei'', honours
Lawrence Morris Lambe Lawrence Morris Lambe (August 27, 1863 – March 12, 1919) was a Canadian geologist, palaeontologist, and ecologist from the Geological Survey of Canada (GSC). His published work, describing the diverse and plentiful dinosaur discoveries from th ...
, the
Canadian Canadians (french: Canadiens) are people identified with the country of Canada. This connection may be residential, legal, historical or cultural. For most Canadians, many (or all) of these connections exist and are collectively the source of ...
geologist A geologist is a scientist who studies the solid, liquid, and gaseous matter that constitutes Earth and other terrestrial planets, as well as the processes that shape them. Geologists usually study geology, earth science, or geophysics, althou ...
and
palaeontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
from the
Geological Survey of Canada The Geological Survey of Canada (GSC; french: Commission géologique du Canada (CGC)) is a Canadian federal government agency responsible for performing geological surveys of the country, developing Canada's natural resources and protecting the en ...
where the holotype was reposited. In 1986 Coombs examined specimen AMNH 5266, at the time by him referred to ''Euoplocephalus'', and determined that it was a juvenile. It consists of five vertebral
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the stores are all owned by individual franchisees. The chain has three different f ...
, a
neural arch 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 characteristic i ...
, one dorsal and two sacral ribs, the right
ischium The ischium () form ...
, the complete right hindlimb, the right pes, an incomplete left pes, and various other fragments. AMNH 5266 was discovered in 1912 at Red Deer River and was collected by Barnum Brown with assistance from Peter Kaisen, George Olsen, and Charles M Sternberg in the sediments from the Horseshoe Canyon Formation.Coombs, W.P., Jr. (1986, June). A Juvenile Ankylosaur Referable to the Genus ''Euoplocephalus'' (Reptilia, Ornithischia). Journal of Vertebrate Paleontology, 6(2), 162-173.


Description

''Anodontosaurus'' was a medium-sized ankylosaurid, measuring up to in length and in body mass. It was a quadrupedal, ground-dwelling herbivore. Like other ankylosaurs, ''Anodontosaurus'' has armor on a majority of the dorsal surfaces of its body. It has a wide, pointed tail club at the end of its armored tail. The skull features postocular caputegulae, which are small polygonal plates of bone that are present on the cranium and are situated to the immediate rear of the eye. Coombs supported the assertion that specimen AMNH 5266 represented a juvenile by citing that the vertebral centra were not fused to their neural arches, and that sacral ribs were likewise not fused to vertebrae and to the ilium. Other morphological characters supporting that this is a juvenile specimen include (a) long bones that feature smooth surfaces, which are not marked by the rugosities characteristic of adult bone; (b) the head of the femur is less spherical in shape and is clearly delimited from the adjacent part of the femoral shaft; (c) the distal ends of the tibia and the fibula are not fused to the astragalus and the calcaneum; and (d) the ungual phalanx of the manus is not widest at the proximal articular end as is observed in adults.


Classification

In 1971,
Walter Coombs Walter may refer to: People * Walter (name), both a surname and a given name * Little Walter, American blues harmonica player Marion Walter Jacobs (1930–1968) * Gunther (wrestler), Austrian professional wrestler and trainer Walter Hahn (born 19 ...
concluded that there was only one species of ankylosaurid during the
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. Campanian s ...
age of the Late Cretaceous of North America. He synonymised the species ''Anodontosaurus lambei'', ''
Dyoplosaurus acutosquameus ''Dyoplosaurus'' (meaning “double-armoured lizard”) is a monospecific genus of ankylosaurid dinosaur from Alberta that lived during the Late Cretaceous (middle Campanian, ~76.5–75 Ma) in what is now the Dinosaur Park Formation. ''Dyoplosa ...
'', and '' Scolosaurus cutleri'' with ''
Euoplocephalus tutus ''Euoplocephalus'' ( ) is a genus of very 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 i ...
''.Coombs W. (1971) The Ankylosauridae. Ph.D. thesis, Columbia University, New York, NY, 487 p. The synonymisation of ''Anodontosaurus lambei'' and ''Euoplocephalus tutus'' was generally accepted and thus CMN 8530 was assigned to ''E. tutus''. However, following the redescription of ''Dyoplosaurus'' as a valid genus by Arbour ''et al.'' (2009), In an abstract presented at the SVP 2010 conference,
Victoria Arbour Victoria Megan Arbour is a Canadian evolutionary biologist and vertebrate palaeontologist at Royal BC Museum, where she is Curator of Palaeontology. An "expert on the armoured dinosaurs known as ankylosaurs", Arbour analyzes fossils and creates ...
considered ''Anodontosaurus'' distinct from ''Euoplocephalus'' in distinctive skull and cervical half ring ornamentation, as well as tail club morphology, including the presence of pointed, triangular knob osteoderms in ''Anodontosaurus''. She therefore reassigned all Horseshoe Canyon Formation ankylosaurine specimens previously referred to ''Euoplocephalus'' to ''Anodontosaurus''. The validity of ''Anodontosaurus'' was formalized in three studies. The first, published by
Paul Penkalski Paul may refer to: *Paul (given name), a given name (includes a list of people with that name) * Paul (surname), a list of people People Christianity *Paul the Apostle (AD c.5–c.64/65), also known as Saul of Tarsus or Saint Paul, early Chri ...
and William T. Blows in 2013, re-validated ''Scolosaurus'' as well. The second study, by Penkalski (2013), named and described ''
Oohkotokia ''Oohkotokia'' ( ) is a genus of ankylosaurid dinosaur within the subfamily Ankylosaurinae. It is known from the upper levels of the Two Medicine Formation (late Campanian stage, about 74 Ma ago) of Montana, United States. The discovery of ''O ...
'' from
Montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columbi ...
on the basis of remains that were originally thought to be referable to ''Euoplocephalus''. Penkalski (2013) performed a small
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis of some ankylosaurine specimens. The only ''Anodontosaurus'' specimen that was included in this analysis was its holotype. ''Anodontosaurus'' was placed in a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with the holotype of ''Euoplocephalus'' and some specimens that are referred to it, while ''Oohkotokia'' was placed in a clade with ''Dyoplosaurus'', and specimens that are thought to represent either ''Dyoplosaurus'' or ''Scolosaurus''. In a study based on the results of her 2010 SVP abstract, Arbour along with Philip Currie formalized the revalidation of ''Anodontosaurus'', and one specimen from the Dinosaur Park Formation, TMP 1997.132.1, was referred to ''Anodontosaurus'' extending the stratigraphic range of the genus back a few million years.Arbour VM, Currie PJ (2013) Euoplocephalus tutus and the Diversity of Ankylosaurid Dinosaurs in the Late Cretaceous of Alberta, Canada, and Montana, USA. PLoS ONE 8(5): e62421. https://doi.org/10.1371/journal.pone.0062421 The DPF specimen, however, was later made the holotype of a new species, ''A. inceptus''.Paul Penkalski (2018). Revised systematics of the armoured dinosaur Euoplocephalus and its allies. Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen 287(3): 261-306. doi: https://doi.org/10.1127/njgpa/2018/0717 The following
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 d ...
is based on a 2015
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
of the Ankylosaurinae conducted by Arbour and Currie:


See also

*
Timeline of ankylosaur research This timeline of ankylosaur research is a chronological listing of events in the history of paleontology focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spikes and sometimes by a club ...
*
2018 in paleontology Flora Plants Fungi Cnidarians Research * New three dimensionally phosphatized microfossils of coronate scyphozoan '' Qinscyphus necopinus'', including a new type of fossil embryo, are described from the Cambrian (Fortunian) Kuanchuanpu F ...


References

{{Taxonbar, from=Q4769855 Ankylosaurids Late Cretaceous dinosaurs of North America Fossil taxa described in 1929 Taxa named by Charles Mortram Sternberg Paleontology in Alberta Campanian genus first appearances Maastrichtian genus extinctions Ornithischian genera