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''Alcovasaurus'' is an extinct genus of herbivorous
stegosauria 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 (North America, Europe a ...
n
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 that lived in 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 ...
. It was found in the
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
of
Natrona County Natrona County is a county in the U.S. state of Wyoming. As of the 2020 United States census, the population was 79,955, making it the second-most populous county in Wyoming. Its county seat is Casper. Natrona County comprises the Casper, WY ...
,
Wyoming Wyoming ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States, Western United States. It borders Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho t ...
, United States. The genus contains a single species, ''Alcovasaurus longispinus'', originally assigned to the genus ''
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 fo ...
''. It is likely a member of the
Dacentrurinae 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 (North America, Europe and Asia), Africa and South A ...
, and has been referred to the genus ''
Miragaia Miragaia () is a former civil parish in the municipality of Porto, Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in contine ...
'' by some authors.


Discovery and naming

In July 1908, Professors
William Harlow Reed William Harlow Reed (9 June 1848 – 24 April 1915) was an American fossil collector and pioneer. He served as a curator at the Museum of Geology at the University of Wyoming, Laramie. He collected for a while for Othniel Charles Marsh but left a ...
and A.C. Dart of the
University of Wyoming The University of Wyoming (UW) is a Public university, public land-grant university, land-grant research university in Laramie, Wyoming, United States. It was founded in March 1886, four years before the territory was admitted as the 44th state, ...
, in the Alcova Quarry in Natrona County, Wyoming, uncovered the skeleton of a stegosaurian. This would be the last major excavation of a dinosaur in which Reed was personally involved. In 1914, the find was named and described as ''Stegosaurus longispinus'' by
Charles Whitney Gilmore Charles Whitney Gilmore (March 11, 1874 â€“ September 27, 1945) was an American paleontologist who gained renown in the early 20th century for his work on vertebrate fossils during his career at the United States National Museum (now the N ...
on the basis of
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 ...
UW 20503 (originally UW D54), a partial postcranial skeleton of an adult individual consisting of forty-two vertebrae, a fragmentary sacrum, two ischia, a portion of one pubis, the right femur, several ribs and four dermal tail spines. 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 ...
is derived from
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 ...
''longus'', "long", and ''spina'', "spine", in reference to the long tail spines. Due to the presence of very long tail spines, ''S. longispinus'' was treated as valid by subsequent authors. In 1993, ''S. longispinus'' was by George Olshevsky and Tracy Lee Ford seen as a possible North American species of the African genus ''
Kentrosaurus ''Kentrosaurus'' ( ; ) is a genus of stegosaurid dinosaur from the Late Jurassic in Lindi Region of Tanzania. The type species is ''K. aethiopicus'', named and described by German people, German Palaeontology, palaeontologist Edwin Hennig in 191 ...
'' as a ?''Kentrosaurus longispinus''. Unfortunately, the type specimen of this species was damaged in the late 1920s when the water pipes of the University of Wyoming's museum burst. For this reason, the whereabouts of the type specimen were mistakenly considered to be lost, although a femur catalogued as part of UW 20503 is still extant, as the last-surviving part of the type specimen. Plaster casts had been made of the rear tail spikes. Also photographic evidence of the skeleton being excavated is still available, showing the bones ''in situ'', as well as of the skeletal museum mount. Although the validity of ''Stegosaurus longispinus'' was disputed because the long dermal spines were likely to be a product of
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the ovum, egg to adult. The term can also be used to refer to t ...
or
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, the amateur freelance paleontologist Roman Ulansky decided that the long tail spines were sufficient to remove ''S. longispinus'' from ''Stegosaurus'' and place it in a new genus, "Natronasaurus". Ulansky interpreted "Natronasaurus" as a close relative of ''Kentrosaurus'' in accordance with the hypothesis of Olshevsky and Ford (1993). Ulansky published the name in a self-published
electronic publication Electronic publishing (also referred to as e-publishing, digital publishing, or online publishing) includes the digital publication of e-books, digital magazines, and the development of digital libraries and catalogues. It also includes the e ...
that was not archived by an independent organization nor had an
ISSN An International Standard Serial Number (ISSN) is an eight-digit to uniquely identify a periodical publication (periodical), such as a magazine. The ISSN is especially helpful in distinguishing between serials with the same title. ISSNs a ...
. This renders the name "Natronasaurus" invalid, and
Peter Malcolm Galton Peter Malcolm Galton (born 14 March 1942 in London) is a British vertebrate paleontologist who has to date written or co-written about 190 papers in scientific journals or chapters in paleontology textbooks, especially on ornithischian and prosau ...
and
Kenneth Carpenter Kenneth Carpenter (born 21 September 1949) is an American 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 are armore ...
gave the genus another name, ''Alcovasaurus'', in 2016, as part of a revision of the species. The new genus name is a reference to Alcova, Wyoming. Galton and Carpenter also referred a very large round spike base to the species, found by Cliff Miles in Wyoming in the 1990s. Its present location is unknown but a cast was made with inventory number DMNH 33431. In 2019, Francisco Costa and
Octávio Mateus Octávio Mateus (born 1975) is a Portugal, Portuguese dinosaur paleontologist and biologist Professor of Paleontology at the Faculdade de Ciências e Tecnologia da NOVA University Lisbon, Universidade Nova de Lisboa. He graduated in University of à ...
judged that ''Alcovasaurus'' was not sufficiently distinct from the genus ''
Miragaia Miragaia () is a former civil parish in the municipality of Porto, Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in contine ...
'', and renamed it ''Miragaia longispinus''. This was based on a description of a new specimen, MG 4863, of ''Miragaia longicollum'' preserving the first relatively complete tail known from a dacentrurine stegosaur. It was concluded that almost all of the diagnostic characters that were found to distinguish ''Alcovasaurus'' from other stegosaurs also occurred in MG 4863. Costa and Mateus considered the two species to be congeneric, with the name ''Miragaia'' having priority. However, a new specimen of ''Dacentrurus'' reported in 2024 from the
Villar del Arzobispo Formation The Villar del Arzobispo Formation is a Late Jurassic to possibly Early Cretaceous geologic formation in eastern Spain. It is equivalent in age to the Lourinhã Formation of Portugal. It was originally thought to date from the Late Tithonian-Mid ...
suggests that ''Alcovasaurus'' is a separate genus and that ''Miragaia'' is a subjective junior synonym of ''Dacentrurus''.


Description

''Alcovasaurus'' was a large stegosaur, reaching in length and in body mass. Some large individuals may have reached in body mass. The thighbone length was by Gilmore determined at . The longest spike was long. Its point was broken and Gilmore estimated the original length of the bone core at . Gilmore first diagnosed ''S. longispinus'' from other ''Stegosaurus'' species by the presence of very long dermal spikes, distal caudal vertebral centra rounded in anterior/posterior view, vestigial transverse processes on distal caudal vertebrae, and centra with mushroom-shaped dorsal extensions. In 2016, Galton and Carpenter indicated five
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...
, unique derived traits. Side processes are present in the distal, rear, and tail vertebrae. The distal tail vertebrae are short, and taller than long. Two tail spike pairs are very slender and elongated, with 90% of the thighbone length. The last spike pair has their greatest width at a quarter of the shaft length, measured from below. The lower joint surface of the condyles of the thighbone is limited to the underside. As a further possible autapomorphy, ''Alcovasaurus'' is different from the Jurassic North American stegosaurians ''Stegosaurus stenops'', ''Stegosaurus sulcatus'' and ''
Hesperosaurus ''Hesperosaurus'' (meaning "western lizard", from Classical Greek (') "western" and (') "lizard") is a herbivorous stegosaurian dinosaur from the Kimmeridgian age of the Jurassic period, approximately 156 million years ago. Fossils of ''Hesper ...
'' in that its sacrum is connected to the ilia by six pairs of sacral ribs instead of four. Other stegosaurians show at most five. Gilmore also claimed that ''Alcovasaurus'' was distinctive in having a rear spike pair equaling the front pair in size. Galton & Carpenter found this trait difficult to confirm from the extant photographic evidence, but it would make ''Alcovasaurus'' differ from all known stegosaurians with four spikes, which always show a shorter rear pair. Also the two pairs would be positioned exceptionally far apart from each other. Galton and Carpenter saw the spikes primarily as weapons. They interpreted the side processes of the caudal vertebrae continuing to the very tail end as a support for an increased muscle mass to swing the tail. This might also be connected to the shortening of the caudal vertebrae, resulting in a tail that was about a quarter shorter than in ''Stegosaurus''. A shorter tail could counteract the
torsion Torsion may refer to: Science * Torsion (mechanics), the twisting of an object due to an applied torque * Torsion of spacetime, the field used in Einstein–Cartan theory and ** Alternatives to general relativity * Torsion angle, in chemistry Bio ...
caused by the greater
moment arm In physics and mechanics, torque is the rotational analogue of linear force. It is also referred to as the moment of force (also abbreviated to moment). The symbol for torque is typically \boldsymbol\tau, the lowercase Greek alphabet, Greek let ...
of the very long spikes. However, the 2019 reinterpretation of ''Alcovasaurus'' as a member of the clade
Dacentrurinae 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 (North America, Europe and Asia), Africa and South A ...
also prompted a reinterpretation of the tail anatomy. Rather than having a strongly muscled tail tip, ''Alcovasaurus'' more likely had very small tail vertebrae past the first third of the tail. Like other dacentrurines, the tail tip probably lacked strong muscles or indeed much soft tissue at all, creating a very flexible, whip-like tail that could be propelled by the heavily muscled tail base.


Classification

Galton and Carpenter considered ''Alcovasaurus'' a member of the
Stegosauridae 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 (North America, Europe and Asia), Africa and South ...
, not more closely related to ''Kentrosaurus'' than to ''Stegosaurus''. A 2017 phylogeny of stegosaurids by Thomas Raven and Susannah Maidment found that ''Alcovasaurus'' lacked the fusion between the trochanters of the femur seen in adult
eurypoda 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 ...
ns (stegosaurians and ankylosaurians), precluding a confident placement as a stegosaur. However, this may be due to the specimen's immaturity, which is impossible to verify since the holotype has been lost. In 2019, Costa and Mateus reinterpreted ''Alcovasaurus'' in the context of newly recognized specimens of ''
Miragaia Miragaia () is a former civil parish in the municipality of Porto, Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in contine ...
'' found in Portugal. These remains allowed them to recognize ''Alcovasaurus'' as a close relative of ''Miragaia'', within the stegosaurid clade
Dacentrurinae 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 (North America, Europe and Asia), Africa and South A ...
. Because ''Alcovasaurus'' shared most of its traits with either ''Dacentrurus'' or ''Miragaia'', Costa and Mateus concluded that it would by definition belong to one of them; they chose ''Miragaia'' to which it was most similar. It remained a separate species though, for which they coined the newly combined name ''Miragaia longispinus''. This combination has not been followed by later research. A 2025 publication by Sánchez-Fenollosa & Cobos included ''Alcovasaurus'' in a
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
and recovered it as the earliest-diverging dacentrurine within the Stegosauridae. These results are displayed in the
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
below:


See also

*
Timeline of stegosaur research This timeline of stegosaur research is a chronological listing of events in the History of paleontology, history of paleontology focused on the stegosaurs, the iconic plate-backed, spike-tailed herbivorous eurypod dinosaurs that predominated duri ...


References

{{Taxonbar, from=Q18684887 Stegosauria Dinosaur genera Kimmeridgian dinosaurs Morrison Formation Dinosaurs of the United States Fossil taxa described in 2016 Taxa named by Peter Galton Taxa named by Kenneth Carpenter