Nodosauridae is a family of

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 ...

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 known 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 ...

to the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...

periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

clade 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

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 ...

, some analyses recover it as a

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

grade leading to the ankylosaurids.

Description

Nodosaurids, like their sister group the ankylosaurids, were heavily armored dinosaurs adorned with rows of bony armor nodules and spines (

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, ...

), which were covered in keratin sheaths. Ankylosaurians were small- to large-sized, heavily built,

quadruped Quadrupedalism is a form of locomotion in which animals have four legs that are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four legs is said to be a quadruped (fr ...

al,

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 ...

s, possessing small, leaf-shaped teeth. Unlike ankylosaurids, nodosaurids lacked mace-like tail clubs and instead had more flexible tail tips. Many nodosaurids had spikes projecting outward from their shoulders. One particularly well-preserved nodosaurid "mummy", the holotype of '' Borealopelta markmitchelli'', preserves a nearly complete set of armor in life position, as well as the keratin covering and mineralized remains of the underlying skin, which indicate reddish pigments in a

countershading Countershading, or Thayer's law, is a method of camouflage in which animal coloration, an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptile ...

pattern.

Classification

The

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

Nodosauridae was erected by

Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of paleontology. A prolific fossil collector, Marsh was one of the preeminent paleontologists of the nineteenth century. Among his legacies are the discovery or ...

in 1890, and anchored on the genus '' Nodosaurus''. The

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 ...

Nodosauridae was first informally defined 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 ankylosaurs closer to ''Panoplosaurus'' than to ''

Ankylosaurus ''Ankylosaurus'' is a genus of Thyreophora, armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period (geology), Period, about 68–66 million years ago, in western North America, m ...

''," a definition followed by Vickaryous,

Teresa Maryańska Teresa Maryańska (1937 – 3 October 2019) was a Polish paleontologist who specialized in Mongolian dinosaurs, particularly pachycephalosaurians and ankylosaurians. She is considered not only as one of Poland's but also one of the world's leadin ...

, and Weishampel in 2004. Vickaryous et al. considered two

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

of nodosaurids to be of uncertain placement (''incertae sedis''): ''Struthiosaurus'' and ''Animantarx'', and considered the most primitive member of the Nodosauridae to be ''Cedarpelta''. Following the publication 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 ...

, Nodosauridae needed to be formally defined following certain parameters, including that the type genus '' Nodosaurus'' was required as an internal specifier. In formally defining Nodosauridae, Madzia and colleagues followed the previously established use for the clade, defining it as the largest clade including '' Nodosaurus textilis'' but not '' Ankylosaurus magniventris''. As all phylogenies referenced included both ''Panoplosaurus'' and ''Nodosaurus'' within the same group relative to ''Ankylosaurus'', the addition of another internal specifier was deemed unnecessary. Nodosauridae is traditionally composed of the basal clade Polacanthinae (sometimes recovered outside of the Nodosauridae), as well as the Panoplosaurini and Struthiosaurini within the Nodosaurinae. Nodosaurinae is defined in the ''

'' as "the largest clade containing '' Nodosaurus textilis'', but not '' Hylaeosaurus armatus'', '' Mymoorapelta maysi'', and '' Polacanthus foxii''". Panoplosaurini is defined in the ''PhyloCode'' as "the largest clade containing '' Panoplosaurus mirus'', but not '' Nodosaurus textilis'' and '' Struthiosaurus austriacus''" while Struthiosaurini has a similar definition of "the largest clade containing '' Struthiosaurus austriacus'', but not '' Nodosaurus textilis'' and '' Panoplosaurus mirus''". Topology A below demonstrates these relationships, following the

phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...

of Rivera-Sylva and colleagues (2018), with clade names added by definition from Madzia et al. (2021). However, in 2023, Raven and colleagues proposed an alternate phylogeny for nodosaurids; instead of the traditional dichotomous split between nodosaurids and ankylosaurids, their analyses resulted in a

grade of these taxa comprising the monophyletic clades Panoplosauridae, Polacanthidae and Struthiosauridae. These results are displayed in Topology B below. Corresponding clades are shown in matching colors for clarity, and ⊞ buttons can be clicked to expand nodes: Nodosaurinae is defined as the largest clade containing '' Nodosaurus textilis'' but not '' Hylaeosaurus armatus'', '' Mymoorapelta maysi'', or '' Polacanthus foxii'', and was formally defined in

2021 Like the year 2020, 2021 was also heavily defined by the COVID-19 pandemic, due to the emergence of multiple Variants of SARS-CoV-2, COVID-19 variants. The major global rollout of COVID-19 vaccines, which began at the end of 2020, continued ...

by Madzia and colleagues, who utilized the name of

Othenio Abel Othenio Lothar Franz Anton Louis Abel (20 June 1875 – 4 July 1946) was an Austrian paleontologist and evolutionary biologist. Together with Louis Dollo, he was the founder of " paleobiology" and studied the life and environment of fossilized ...

1919 Events January * January 1 ** The Czechoslovak Legions occupy much of the self-proclaimed "free city" of Pressburg (later Bratislava), enforcing its incorporation into the new republic of Czechoslovakia. ** HMY ''Iolaire'' sinks off th ...

, who created the term to unite ''

'', ''

Hierosaurus ''Hierosaurus'' (meaning "sacred lizard") is an extinct genus of Nodosauridae, nodosaurid ankylosaur which lived during the Late Cretaceous 87 to 82 million years ago. Its fossils were found in the Smoky Hill Chalk, Smoky Hill Chalk Member of the ...

'' and '' Stegopelta''. The name has been significantly refined in content since Abel first used it to unite all quadrupedal, plate-armoured ornithischians, now including a significant number of taxa from the

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 ...

through

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

of Europe, North America, and

Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...

. Previous informal definitions of the group described the clade as all taxa closer to '' Panoplosaurus'', or ''Panoplosaurus'' and ''Nodosaurus'', than to the early ankylosaurs '' Sarcolestes'', ''Hylaeosaurus'', ''Mymoorapelta'' or ''Polacanthus'', which was reflected in the specifiers chosen by Madzia et al. when formalizing the clade following the

. The

2018 Events January * January 1 – Bulgaria takes over the Presidency of the Council of the European Union, after the Estonian presidency. * January 4 – SPLM-IO rebels loyal to Chan Garang Lual start a raid against Juba, capital of ...

phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Panoplosaurini by Madzia et al., in addition to the supplemental analyses of Thompson et al. (2012), Arbour and Currie (2016), Arbour et al. (2016), and Brown et al. (2017). Panoplosaurini is defined as the largest clade containing '' Panoplosaurus mirus'', but not '' Nodosaurus textilis'' or '' Struthiosaurus austriacus'', and was named in

by Madzia and colleagues for the group found in many previous analyses, both morphological and phylogenetic. Panoplosaurini includes not only the

''Panoplosaurus'', '' Denversaurus'' and '' Edmontonia'', but also the mid

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 ...

Animantarx ''Animantarx'' ( ; meaning 'living citadel') is a genus of nodosaurid ankylosaurian dinosaur from the Early Cretaceous, Early and Late Cretaceous of western North America. Like other nodosaurs, it would have been a slow-moving quadrupedal herbivo ...

'' and '' Texasetes'', as well as ''

Patagopelta ''Patagopelta'' (meaning "Patagonian shield") is an extinct genus of ankylosaurian dinosaur from the Late Cretaceous (upper Campanian–lower Maastrichtian) Allen Formation of Argentina. The genus contains a Monotypic taxon, single species, ''P. ...

''. However, in the study describing it, its authors only placed it as a nodosaurine outside Panoplosaurini. The approximately equivalent clade Panoplosaurinae, named in

1929 This year marked the end of a period known in American history as the Roaring Twenties after the Wall Street Crash of 1929 ushered in a worldwide Great Depression. In the Americas, an agreement was brokered to end the Cristero War, a Catholic ...

Franz Nopcsa Franz may refer to: People * Franz (given name) * Franz (surname) Places * Franz (crater), a lunar crater * Franz, Ontario, a railway junction and unorganized town in Canada * Franz Lake, in the state of Washington, United States – see Fran ...

, but was not significantly used until

Robert Bakker Robert Thomas Bakker (born March 24, 1945) is an American paleontologist who helped reshape modern theories about dinosaurs, particularly by adding support to the theory that some dinosaurs were endothermic (warm-blooded). Along with his mentor ...

reused the name in 1988, alongside the new clades Edmontoniinae and Edmontoniidae, which were considered to unite ''Panoplosaurus'', ''Denversaurus'' and ''Edmontonia'' to the exclusion of other ankylosaurs. As none of the clades were commonly used, or formally named following the

, Madzia et al. named Panoplosaurini instead, as the group of taxa fell within the clade Nodosaurinae, and having the same ''-inae'' suffix on both parent and child taxon could be confusing in future. The

phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Panoplosaurini by Madzia et al., in addition to the supplemental analyses of Arbour et al. (2016), Brown et al. (2017), and Zheng et al. (2018). Struthiosaurini is defined as the largest clade containing '' Struthiosaurus austriacus'', but not '' Nodosaurus textilis'' or '' Panoplosaurus mirus'', and was named in

by Madzia and colleagues for the relatively stable group found in many previous analyses. Struthiosaurini includes not only the

European ''Struthiosaurus'', but also the

European ''

Europelta ''Europelta'' (meaning “Europe’s shield”) is a monospecific genus of nodosauridae, nodosaurid dinosaur from Spain that lived during the Early Cretaceous (early Albian stage, ~113.0 Ma) in what is now the lower Escucha Formation of the Terue ...

'', the Late Cretaceous European '' Hungarosaurus'', and '' Stegopelta'' and '' Pawpawsaurus'' from the mid Cretaceous of North America. The approximately equivalent clade Struthiosaurinae, named in

1923 In Greece, this year contained only 352 days as 13 days was skipped to achieve the calendrical switch from Julian to Gregorian Calendar. It happened there that Wednesday, 15 February ''(Julian Calendar)'' was followed by Thursday, 1 March ' ...

was previously used to include European nodosaurids, but was never formally named following the

, so Madzia et al. named Struthiosaurini instead, as the group of taxa fell within the clade Nodosaurinae, and having the same ''-inae'' suffix on both parent and child taxon could be confusing in future. The

phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Struthiosaurini by Madzia et al., in addition to the supplemental analyses of Arbour et al. (2016), Brown et al. (2017), and Zheng et al. (2018).

Biogeography

Nodosaurids are known from diverse remains throughout Europe, Asia, and North America. Some

Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...

n ankylosaurs, including the Antarctican ''

Antarctopelta ''Antarctopelta'' (; meaning 'Antarctic shield') is a genus of ankylosaurian dinosaur, a group of large, quadrupedal herbivores, that lived during the Maastrichtian stage of the Late Cretaceous period on what is now James Ross Island, Antarctica ...

'' and Argentinian ''

'', were originally regarded as belonging to the Nodosauridae, but later analyses provided support for them belonging to the

Parankylosauria Parankylosauria is a group of Basal (phylogenetics), 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 ...

, a separate lineage of basal ankylosaurs restricted to the Southern Hemisphere. ImageSize = width:1250px height:auto barincrement:15px PlotArea = left:10px bottom:50px top:10px right:10px Period = from:-175.6 till:-55.8 TimeAxis = orientation:horizontal ScaleMajor = unit:year increment:10 start:-170 ScaleMinor = unit:year increment:1 start:-175 TimeAxis = orientation:hor AlignBars = justify Colors = #legends id:triassic value:rgb(0.51,0.17,0.57) id:jurassic value:rgb(0.2,0.7,0.79) id:earlyjurassic value:rgb(0,0.69,0.89) id:middlejurassic value:rgb(0.52,0.81,0.91) id:latejurassic value:rgb(0.74,0.89,0.97) id:cretaceous value:rgb(0.5,0.78,0.31) id:earlycretaceous value:rgb(0.63,0.78,0.65) id:latecretaceous value:rgb(0.74,0.82,0.37) id:paleogene value:rgb(0.99,0.6,0.32) id:paleocene value:rgb(0.99,0.65,0.37) BarData= bar:eratop bar:space bar:periodtop bar:space bar:NAM1 bar:NAM2 bar:NAM3 bar:NAM4 bar:NAM5 bar:NAM6 bar:NAM7 bar:NAM8 bar:NAM9 bar:NAM10 bar:NAM11 bar:NAM12 bar:NAM13 bar:NAM14 bar:NAM15 bar:NAM16 bar:NAM17 bar:NAM18 bar:NAM19 bar:NAM20 bar:NAM21 bar:NAM22 bar:NAM23 bar:space bar:period bar:space bar:era PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 shift:(7,-4) bar:eratop from: -175.6 till: -145.5 color:jurassic text:

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. ...

from: -145.5 till: -66.0 color:cretaceous text:

from: -66.0 till: -55.8 color:paleogene text:

Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...

bar:periodtop from: -175.6 till: -161.2 color:middlejurassic text:

Middle Middle or The Middle may refer to: * Centre (geometry), the point equally distant from the outer limits. Places * Middle (sheading), a subdivision of the Isle of Man * Middle Bay (disambiguation) * Middle Brook (disambiguation) * Middle Creek ...

from: -161.2 till: -145.5 color:latejurassic text:

Late Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudonym used by Dave Groh ...

from: -145.5 till: -99.6 color:earlycretaceous text: Early from: -99.6 till: -66.0 color:latecretaceous text:

from: -66.0 till: -55.8 color:paleocene text: Paleo. PlotData= align:left fontsize:M mark:(line,white) width:5 anchor:till align:left color:triassic bar:NAM1 from:-166 till:-164 text: Sarcolestes? color:triassic bar:NAM2 from:-157 till:-153 text: Gargoyleosaurus color:triassic bar:NAM3 from:-157 till:-145 text:

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 ...

? color:triassic bar:NAM4 from:-145 till:-100 text: Taohelong color:triassic bar:NAM5 from:-137 till:-134 text: Hylaeosaurus? color:triassic bar:NAM6 from:-131 till:-126 text:

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 ...

color:triassic bar:NAM7 from:-131 till:-113 text: Sauroplites color:triassic bar:NAM1 from:-128 till:-126 text: Gastonia color:triassic bar:NAM2 from:-128 till:-126 text: Horshamosaurus color:triassic bar:NAM3 from:-128 till:-126 text: Hoplitosaurus color:triassic bar:NAM1 from:-113 till:-94 text: Dongyangopelta color:triassic bar:NAM2 from:-105 till:-100 text: Anoplosaurus color:triassic bar:NAM3 from:-97 till:-95 text: Acanthopholis color:jurassic bar:NAM8 from:-118 till:-108 text: Sauropelta color:jurassic bar:NAM9 from:-118 till:-108 text: Tatankacephalus color:jurassic bar:NAM10 from:-116 till:-113 text: Propanoplosaurus color:jurassic bar:NAM11 from:-113 till:-108 text: Priconodon color:jurassic bar:NAM12 from:-113 till:-100 text: Silvisaurus color:jurassic bar:NAM13 from:-112 till:-107 text: Borealopelta color:jurassic bar:NAM14 from:-100 till:-94 text: Zhejiangosaurus? color:jurassic bar:NAM15 from:-100 till:-94 text: Nodosaurus color:jurassic bar:NAM16 from:-99 till:-96 text: Peloroplites color:jurassic bar:NAM8 from:-86 till:-84 text: Acantholipan color:jurassic bar:NAM9 from:-86 till:-84 text:

Niobrarasaurus ''Niobrarasaurus'' (meaning "Niobrara lizard") is an extinct genus of Nodosauridae, nodosaurid ankylosaur which lived during the Cretaceous 87 to 82 million years ago. Its fossils were found in the Smoky Hill Chalk, Smoky Hill Chalk Member of the ...

color:jurassic bar:NAM10 from:-84 till:-75 text: Danubiosaurus? color:jurassic bar:NAM11 from:-84 till:-75 text: Rhadinosaurus? color:jurassic bar:NAM12 from:-81 till:-79 text: Invictarx color:jurassic bar:NAM13 from:-75 till:-74 text: Ahshislepelta? color:jurassic bar:NAM14 from:-72 till:-70 text: Glyptodontopelta color:cretaceous bar:NAM17 from:-113 till:-108 text:

color:cretaceous bar:NAM18 from:-103 till:-100 text: Pawpawsaurus color:cretaceous bar:NAM19 from:-97 till:-95 text: Stegopelta color:cretaceous bar:NAM17 from:-86 till:-84 text: Hungarosaurus color:cretaceous bar:NAM18 from:-84 till:-66 text: Struthiosaurus color:paleogene bar:NAM20 from:-103 till:-100 text: Texasetes color:paleogene bar:NAM21 from:-99 till:-96 text:

color:paleogene bar:NAM21 from:-77 till:-72 text: Edmontonia color:paleogene bar:NAM22 from:-76 till:-75 text: Panoplosaurus color:paleogene bar:NAM23 from:-68 till:-66 text: Denversaurus PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 bar:period from: -175.6 till: -161.2 color:middlejurassic text:

from: -161.2 till: -145.5 color:latejurassic text:

from: -145.5 till: -99.6 color:earlycretaceous text: Early from: -99.6 till: -66.0 color:latecretaceous text:

from: -66.0 till: -55.8 color:paleocene text: Paleo. bar:era from: -175.6 till: -145.5 color:jurassic text:

from: -145.5 till: -66.0 color:cretaceous text:

from: -66.0 till: -55.8 color:paleogene text:

References

External links

Alberta oilsands discovery of 2011
{{Authority control Dinosaur families

Description

Classification

Biogeography

See also

References

Further reading

External links