''Struthiosaurus'' (
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
''struthio'' = ostrich +
Greek
Greek may refer to:
Greece
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 ...
''sauros'' = lizard) is a
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
nodosaurid
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 ...
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, from the Late
Cretaceous
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 th ...
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 mya. The ...
-
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 ...
) of
Austria
Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
,
Romania
Romania ( ; ro, România ) is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern, and Southeast Europe, Southeastern Europe. It borders Bulgaria to the south, Ukraine to the north, Hungary to the west, S ...
,
France
France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
and
Hungary
Hungary ( hu, Magyarország ) is a landlocked country in Central Europe. Spanning of the 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 a ...
in
Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
.
[''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 t ...
at the ''Gute Hoffnung'' coal mine at
Muthmannsdorf near
Wiener Neustadt
Wiener Neustadt (; ; Central Bavarian: ''Weana Neistod'') 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 administration of Wiener Neustadt-Land Distr ...
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 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 herpetology. He died of ...
. The marl was a
fresh water
Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...
deposit, now considered part of the
Grünbach Formation
The Grünbach Formation is an Austrian geological formation that dates to the lower Campanian age of the Late Cretaceous. it forms part of the Gosau Group, and represents a marine regression event, representing a coastal/brackish environment, bein ...
.
The finds were stored in the museum of the
University of Vienna
The University of Vienna (german: Universität Wien) is a public research university located in Vienna, Austria. It was founded by Duke Rudolph IV in 1365 and is the oldest university in the German-speaking world. With its long and rich histor ...
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 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''; plural 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 nam ...
and gave no
etymology
Etymology ()The New Oxford Dictionary of English (1998) – p. 633 "Etymology /ˌɛtɪˈmɒlədʒi/ the study of the class in words and the way their meanings have changed throughout time". is the study of the history of the Phonological chan ...
of the name. The generic name is derived from
new Latin
New Latin (also called Neo-Latin or Modern Latin) is the revival of Literary Latin used in original, scholarly, and scientific works since about 1500. Modern scholarly and technical nomenclature, such as in zoological and botanical taxonomy ...
''struthio'', itself derived from
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 ...
στρούθειος, ''stroutheios'', "of the ostrich". Bunzel chose the name because of the birdlike morphology of the braincase.
The
specific name 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 gen ...
'' 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 in ...
'' 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 body ...
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 (medicine), nodule, small wikt:eminence, eminence, or warty outgrowth found on external or internal Organ (anatomy), organs of a plant or an animal.
In plant ...
, 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") is the type genus (defining example) of the mosasaurs, an extinct group of aquatic squamate reptiles. It lived from about 82 to 66 million years ago during the Campanian and Maastrichtian stages o ...
'', the giant lizard named after the
River Maas
The Meuse ( , , , ; wa, Moûze ) or Maas ( , ; li, Maos or ) 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 t ...
, he named this lizard ''Danubiosaurus anceps'', after the
Danube
The Danube ( ; ) is a river that was once a long-standing frontier of the Roman Empire and today connects 10 European countries, running through their territories or being a border. Originating in Germany, the Danube flows southeast for , pa ...
. The specific name ''anceps'' means "double-headed" in
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
, 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
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 ...
have been referred to ''Struthiosaurus'', most based on very fragmentary and nondiagnostic material. Three valid species are recognized by
paleontologist
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 ...
s: ''S. austriacus'' Bunzel, 1871, 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 ...
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
Nodosaurinae is a subfamily of nodosaurid ankylosaurs from the Cretaceous of Europe, North America, and South America. The group is defined as the largest clade containing '' Nodosaurus textilis'' but not ''Hylaeosaurus armatus'', '' Mymoorapelta ...
tribe
Struthiosaurini
Struthiosaurini (derived from ''Struthiosaurus'', "ostrich reptile") is a clade of nodosauridae, nodosaurid ankylosaurs from the Cretaceous of Europe and North America. The group is defined as the largest clade containing ''Struthiosaurus austriacu ...
, members of which are found only in Europe.
![Struthiosaurus Nopsca 1915](https://upload.wikimedia.org/wikipedia/commons/6/61/Struthiosaurus_Nopsca_1915.png)
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 fat ...
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. Campanian s ...
-
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 ...
-age rocks of southern
France
France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
, 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 s ...
'' and referred to Nodosauridae ''
incertae sedis
' () or ''problematica'' is a term used for a 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 (singular: 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 ...
. 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
* Dorsal co ...
vertebrae. It differs from ''S. austriacus'' in the shape of the
. (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 vertebra.
The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
. 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 overlies the occipital lobes of the cereb ...
. 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 specialising in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution.
The storie ...
in
London
London is the capital and largest city of England and the United Kingdom, with a population of just under 9 million. It stands on the River Thames in south-east England at the head of a estuary down to the North Sea, and has been a majo ...
, 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 Sau ...
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
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, were ...
. He later corrected its name to
Acanthopholidae.
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 ...
in 1978 stated it was a nodosaurid.
Cladistic
Cladistics (; ) 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 typically shared derived char ...
analysis of ''Struthiosaurus'' indicates that the taxon is a member of the
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 suggested it may be one of the most basal ankylosaurs in the
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Ankylosauria
Ankylosauria is a group of herbivorous dinosaurs of the order 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
2018
File:2018 Events Collage.png, From top left, clockwise: The 2018 Winter Olympics opening ceremony in PyeongChang, South Korea; Protests erupt following the Assassination of Jamal Khashoggi; March for Our Lives protests take place across the United ...
phylogenetic analysis of Rivera-Sylva and colleagues, resolving ''Struthiosaurus'' species within the clade
Struthiosaurini
Struthiosaurini (derived from ''Struthiosaurus'', "ostrich reptile") is a clade of nodosauridae, nodosaurid ankylosaurs from the Cretaceous of Europe and North America. The group is defined as the largest clade containing ''Struthiosaurus austriacu ...
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 focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spikes and sometimes by a club ...
* ''
Cairanoolithus''
References
{{Authority control
Nodosaurids
Ornithischian genera
Maastrichtian life
Late Cretaceous dinosaurs of Europe
Cretaceous Austria
Cretaceous France
Cretaceous Romania
Fossils of Austria
Fossils of France
Fossils of Romania
Hațeg fauna
Fossil taxa described in 1871
Taxa named by Emanuel Bunzel