Danubiosaurus
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''Struthiosaurus'' (
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''struthio'' = ostrich +
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
''sauros'' = lizard) is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
nodosaurid Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the s ...
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, from the Late
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 ...
period (
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya ( million years ago) and 83.6 ± 0.7 m ...
-
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
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,
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,
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
and
Hungary Hungary is a landlocked country in Central Europe. Spanning much of the Pannonian Basin, Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia and ...
in
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 ...
.''Struthiosaurus'' in The Dinosaur Encyclopaedia
at Dino Russ's Lair
It was a small dinosaur, measuring in length and weighing .


History of discovery

In 1859, geologist
Eduard Suess Eduard Suess (; 20 August 1831 – 26 April 1914) was an Austrian geologist and an expert on the geography of the Alps. He is responsible for hypothesising two major former geographical features, the supercontinent Gondwana (proposed in 1861) and ...
at the ''Gute Hoffnung'' coal mine at Muthmannsdorf near
Wiener Neustadt Wiener Neustadt (; Lower_Austria.html" ;"title=".e. Lower Austria">.e. Lower Austria , ) is a city located south of Vienna, in the state of Lower Austria, in northeast Austria. It is a self-governed city and the seat of the district administr ...
in Austria, discovered a dinosaur tooth on a stone pile. With the help of mine intendant Pawlowitsch it was attempted to find the source of the fossil material. The search proved fruitless at first but ultimately a thin marl layer was discovered, intersected by an obliquely sloping mine shaft, which contained an abundant number of various bones. These were subsequently excavated by Suess and
Ferdinand Stoliczka Ferdinand Stoliczka (Czech language, Czech written Stolička, 7 June 1838 – 19 June 1874) was a Moravian palaeontologist who worked in India on paleontology, geology and various aspects of zoology, including ornithology, malacology, and herpe ...
. The marl was a
fresh water Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salt (chemistry), salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include ...
deposit, now considered part of the
Grünbach Formation The Grünbach Formation is an Austrian geological formation that dates to the early Campanian age of the Late Cretaceous. it forms part of the Gosau Group, and represents a marine regression event, representing a coastal/brackish environment, bei ...
. The finds were stored in the museum of the
University of Vienna The University of Vienna (, ) is a public university, public research university in Vienna, Austria. Founded by Rudolf IV, Duke of Austria, Duke Rudolph IV in 1365, it is the oldest university in the German-speaking world and among the largest ...
but received little attention until they were studied by
Emanuel Bunzel Emanuel Bunzel (1828–1895), was an Austrian paleontologist. Biography Emanuel Bunzel was born in Prague, Kingdom of Bohemia (today the Czech Republic), in 1828. In 1871, he described a skull fragment found in an Austrian coal mine in 1859 by ...
in 1870. In 1871, Bunzel published a treatise describing the fossils and naming several new genera and species. One of them was the genus ''Struthiosaurus'' based on a single partial portion of the posterior end of the skull, largely consisting of the braincase. The
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...
and only known species of the genus at the time was ''Struthiosaurus austriacus''. Bunzel stated that he only provisionally named the
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
and gave no
etymology Etymology ( ) is the study of the origin and evolution of words—including their constituent units of sound and meaning—across time. In the 21st century a subfield within linguistics, etymology has become a more rigorously scientific study. ...
of the name. The generic name is derived from
Neo-Latin Neo-LatinSidwell, Keith ''Classical Latin-Medieval Latin-Neo Latin'' in ; others, throughout. (also known as New Latin and Modern Latin) is the style of written Latin used in original literary, scholarly, and scientific works, first in Italy d ...
''struthio'', itself derived from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
στρούθειος, ''stroutheios'', "of the ostrich". Bunzel chose the name because of the birdlike morphology of the braincase. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
refers to the provenance from Austria. Apart from the braincase, Bunzel unknowingly described other material of ''Struthiosaurus''. He recognized that there were bones and osteoderms of armoured dinosaurs among the finds and referred them to a ''
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 ge ...
'' sp. and a ''
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 ...
'' sp. These British genera represented the best known
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 bod ...
n forms found at the time. Bunzel also discovered two rib fragments which had a very puzzling build. They were double-headed but the upper rib head, the
tuberculum In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection, b ...
, was short and positioned in such a way that it could not possibly touch the vertebra, if the shaft was oriented in the usual vertical position. He assumed that only the lower capitulum connected to the vertebral body. A rib touching the vertebra with a single surface is normal for lizards, though in their case the rib heads are fused into a single synapophysis. Bunzel therefore concluded that the ribs belonged to a giant lizard. In analogy to ''
Mosasaurus ''Mosasaurus'' (; "lizard of the Meuse (river), Meuse River") is the type genus (defining example) of the mosasaurs, an extinct group of aquatic Squamata, squamate reptiles. It lived from about 82 to 66 million years ago during the Campanian an ...
'', the giant lizard named after the
River Maas The Meuse or Maas is a major European river, rising in France and flowing through Belgium and the Netherlands before draining into the North Sea from the Rhine–Meuse–Scheldt delta. It has a total length of . History From 1301, the upper ...
, he named this lizard ''Danubiosaurus anceps'', after the
Danube The Danube ( ; see also #Names and etymology, other names) is the List of rivers of Europe#Longest rivers, second-longest river in Europe, after the Volga in Russia. It flows through Central and Southeastern Europe, from the Black Forest sou ...
. The specific name ''anceps'' means "double-headed" in
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
, highlighting the, for a lizard, exceptional trait of having double-headed ribs. In fact the ribs were those of ''Struthiosaurus''. In Ankylosauria, the rump is so flat that the upper part of the rib shafts sticks out sideways, which rotates the short tuberculum to the diapophysis, its vertebral contact facet. Many
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
have been referred to ''Struthiosaurus'', most based on very fragmentary and nondiagnostic material. Three valid species are recognized by
paleontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
s: ''S. austriacus'' Bunzel, 1871, based on
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
PIWU 2349/6; ''S. transylvanicus'' Nopcsa, 1915, based on BMNH R4966, a skull and partial skeleton from Romania; and ''S. languedocensis'' Garcia and Pereda-Suberbiola, 2003, based on UM2 OLV-D50 A–G CV, a partial skeleton found in 1998 in France. It is the namesake of the nodosaurine tribe
Struthiosaurini Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the s ...
, members of which are found only in Europe. A number of invalid taxa have been shown to be junior synonyms of ''Struthiosaurus austriacus'', most of them created when
Harry Govier Seeley Harry Govier Seeley (18 February 1839 – 8 January 1909) was a British paleontologist. Early life Seeley was born in London on 18 February 1839, the second son of Richard Hovill Seeley, a goldsmith, and his second wife Mary Govier. When his fa ...
in 1881 revised the Austrian material. They include: ''Danubiosaurus anceps'' Bunzel, 1871; ''Crataeomus pawlowitschii'' Seeley, 1881; ''Crataeomus lepidophorus'' Seeley 1881; ''Pleuropeltis suessii'' Seeley, 1881; ''Rhadinosaurus alcimus'' Seeley 1881, ''Hoplosaurus ischyrus'' Seeley 1881 and ''Leipsanosaurus noricus'' Nopcsa, 1918. Another European ankylosaurid, ''Rhodanosaurus ludguensis'' Nopcsa, 1929, from
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. Campa ...
-
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 ...
-age rocks of southern
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
, is now regarded as a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
'' and referred to Nodosauridae ''
incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
''. The three valid species of ''Struthiosaurus'' differ from one another in that ''S. austriacus'' is smaller than ''S. transylvanicus'' and possesses less elongate
cervical vertebrae In tetrapods, cervical vertebrae (: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In saurop ...
. Also, though the quadrate-paroccipital process contact is fused in ''S. transylvanicus'', it is unfused in ''S. austriacus''. The skull of ''S. languedocensis'' is unknown, but the taxon differs from ''S. transylvanicus'' in the flatter shape of the
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...
vertebrae. It differs from ''S. austriacus'' in the shape of the
ischium The ischium (; : is ...
. (Vickaryous, Maryanska, and Weishampel 2004)


Classification

Bunzel was very puzzled by the braincase. He knew that it belonged to a reptile instead of a mammal because of a single as opposed to a double-headed
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the Atlas (anatomy), atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape ...
. The back of the head was otherwise not very reptilian as it was low, compact, fused and convex in a gradual curve towards the skull-roof. Lizards had a very different, more "open",
occiput The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone lies over the occipital lobes of the ...
. Crocodiles were more similar but still had a concave skull rear. Bunzel considered whether it might be a dinosaur but in 1871 little dinosaurian occiput material had been described and it seemed to him that their skulls in this respect were more lizard-like. The only group showing a comparable rounding and fusion of skull bones were the birds. Bunzel sent a drawing and description to Professor
Thomas Huxley Thomas Henry Huxley (4 May 1825 – 29 June 1895) was an English biologist and anthropologist who specialized in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The stor ...
in
London London is the Capital city, capital and List of urban areas in the United Kingdom, largest city of both England and the United Kingdom, with a population of in . London metropolitan area, Its wider metropolitan area is the largest in Wester ...
, at the time one of the few dinosaur experts. Huxley agreed that the braincase resembled that of a bird, commenting "This skull-fragment is more bird-like, than any thing icI have yet seen". Knowing that Huxley had named a reptile order
Ornithoscelida Ornithoscelida () is a proposed clade that includes various major groupings of dinosaurs. An order Ornithoscelida was originally proposed by Thomas Henry Huxley but later abandoned in favor of Harry Govier Seeley's division of Dinosauria into ...
for forms sharing with birds certain traits in the pelvis and hindlimbs, Bunzel ended his description with the prediction that "with time, it might also be possible to create an order ''Ornithocephala'' ('Bird Heads')". Bunzel was correct in assuming an affinity with birds but this was because birds are themselves dinosaurs. In dinosaurs, the rear skull bones are generally strongly fused. Nodosaurids did convergently develop a rounded skull. As the massive quadrates were lacking, the skull fragment gave a false impression of being lightly built. Ankylosaur material at the time was typically referred to the
Scelidosauridae ''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 ...
but because this was the first ankylosaur braincase to be described, the connection was not obvious. The first to understand it represented an armoured dinosaur was Nopcsa who in 1902 placed it in the Acanthopholididae. He later corrected its name to Acanthopholidae. Walter Coombs in 1978 stated it was a nodosaurid.
Cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analysis of ''Struthiosaurus'' indicates that the taxon is a member of the
Nodosauridae Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the ...
and suggested it may be one of the most basal ankylosaurs in 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 ...
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 ...
. An analysis by Ösi in 2005, describing the taxon ''Hungarosaurus'', found that while being younger in age than other nodosaurids, ''Struthiosaurus'' was one of the more basal taxa, although many features could not be coded for it. The cladogram below shows the results of the
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phylogenetic analysis of Rivera-Sylva and colleagues, resolving ''Struthiosaurus'' species within the clade
Struthiosaurini Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the s ...
as labelled by Madzia ''et al.'', with relationships outside Struthiosaurini excluded for simplicity.


Paleobiology

Examinations of the brain of ''Struthiosaurus'' published in 2022 suggest it had very poor hearing and relied mostly on its armor for defense against predators. Its flocculus was very small and its lagena were very short, suggesting it was very sluggish in nature as well as solitary.


See also

*
Timeline of ankylosaur research This timeline of ankylosaur research is a chronological listing of events in the History of paleontology, history of paleontology focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spik ...
* '' Cairanoolithus''


References

{{Authority control Nodosauridae Dinosaur genera Late Cretaceous dinosaurs Hațeg fauna Taxa named by Emanuel Bunzel Fossil taxa described in 1871 Dinosaurs of France Dinosaurs of Romania Dinosaurs of Europe