Nanosaurus

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Nanosaurus
Temporal range: Late Jurassic, 155–148  Ma
Othnielosaurus fruita 1.jpg
Reconstructed skeleton cast, Dinosaur Journey Museum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Ornithischia
Clade: Genasauria
Clade: Neornithischia
Family: Nanosauridae
Marsh, 1877
Genus: Nanosaurus
Marsh, 1877
Species:
N. agilis
Binomial name
Nanosaurus agilis
Marsh, 1877
Synonyms
Genus synonymy
  • DrinkerBakker et al., 1990
  • OthnieliaGalton, 1977
  • OthnielosaurusGalton, 2007
Species synonymy
  • Drinker nistiBakker et al., 1990
  • Laosaurus consorsMarsh, 1894
  • Nanosaurus rexMarsh, 1877
  • Othnielia rex(Marsh, 1877)
  • Othnielosaurus consors(Marsh, 1894)

Nanosaurus ("small or dwarf lizard") is the name given to a genus of neornithischian dinosaur that lived about 155 to 148 million years ago, during the Late Jurassic-age. Its fossils are known from the Morrison Formation of the south-western United States. The type and only species, Nanosaurus agilis, was described and named by Othniel Charles Marsh in 1877. The taxon has a complicated taxonomic history, largely the work of Marsh and Peter M. Galton, involving the genera Laosaurus , Hallopus , Drinker, Othnielia, and Othnielosaurus, the latter three now being considered to be synonyms of Nanosaurus. It had historically been classified as a hypsilophodont or fabrosaur, types of generalized small bipedal herbivore, but more recent research has abandoned these groupings as paraphyletic and Nanosaurus is today considered a basal member of Neornithischia.

Contents

Description

Size compared to a human Othnielosaurus Size Comparison by PaleoGeek.svg
Size compared to a human

Nanosaurus is known from material from all parts of the body, including two good skeletons, although the skull is still poorly known. [1] It was a small animal, with specimens previously assigned to Drinker and Othnielosaurus measuring 2–2.2 metres (6.6–7.2 ft) long and weighing 20–30 kilograms (44–66 lb). [2]

It was a bipedal dinosaur with short forelimbs and long hindlimbs with large processes for muscle attachments. [3] The hands were short and broad with short fingers. The head was small. It had small leaf-shaped cheek teeth (triangular and with small ridges and denticles lining the front and back edges), and premaxillary teeth with less ornamentation. [4]

Like several other neornithischian dinosaurs, such as Hypsilophodon , Thescelosaurus , and Talenkauen , Nanosaurus had thin plates lying along the ribs. Called intercostal plates, these structures were cartilaginous in origin. [5]

History and taxonomy

Marsh's original groundwork

Holotype dentary and ilium Nanosaurus.jpg
Holotype dentary and ilium
Type specimen of Nanosaurus agilis Marsh 1877 (YPM VP 1913) as illustrated in 1908 (left) and cast of the bone impressions after the fragmentary bone was removed (right) Nano type.jpg
Type specimen of Nanosaurus agilis Marsh 1877 (YPM VP 1913) as illustrated in 1908 (left) and cast of the bone impressions after the fragmentary bone was removed (right)

Nanosaurus has had a long and complicated taxonomic history. In 1877, Marsh named two species of Nanosaurus in separate publications, based on partial remains from the Morrison Formation of Garden Park, Colorado. One paper described N. agilis, based on YPM 1913, with remains including impressions of a dentary, and postcranial bits including an ilium, thigh bones, shin bones, and a fibula. [6] The other paper named N. rex, a second species which Marsh based on YPM 1915 (also called 1925 in Galton, 2007), a complete thigh bone. [4] [7] He regarded both species as small ("fox-sized") animals. [7] A third species, N. victor , was named, which he soon recognized to be something completely different, and is now known as the small, bipedal crocodylomorph Hallopus . [6] [8]

Othniel Charles Marsh's 1896 skeletal restoration of "Laosaurus" consors (now Nanosaurus). Marsh laosaurus.jpg
Othniel Charles Marsh's 1896 skeletal restoration of "Laosaurus" consors (now Nanosaurus).

The next year, he named the new genus Laosaurus on material collected by Samuel Wendell Williston from Como Bluff, Wyoming. Two species were named: the type species L. celer, based on parts of eleven vertebrae (YPM 1875); [9] and the "smaller" L. gracilis, originally based on a back vertebra's centrum, a caudal centrum, and part of an ulna (review by Peter Galton in 1983 finds the specimen to now consist of thirteen back and eight caudal centra, and portions of both hindlimbs). [9] [10] A third species, L. consors, was established by Marsh in 1894 for YPM 1882, which consists of most of one articulated skeleton and part of at least one other individual. [11] The skull was only partially preserved, and the fact that the vertebrae were represented only by centra suggests a partially grown individual. Galton (1983) notes that much of the current mounted skeleton was restored in plaster, or had paint applied. [10]

Galton's taxonomic revisions

Cast mounted as if a herd running, Denver Museum of Nature and Science. Othnielia.png
Cast mounted as if a herd running, Denver Museum of Nature and Science.
Skull-less skeleton of a juvenile Nanosaurus skeleton (BYU 163) Nanosaurus BYU 163.tif
Skull-less skeleton of a juvenile Nanosaurus skeleton (BYU 163)

These animals attracted little professional attention until the 1970s and 1980s, when Peter Galton reviewed many of the "hypsilophodonts" in a series of papers. In 1973, he and Jim Jensen described a partial skeleton (BYU ESM 163 as of Galton, 2007) missing the head, hands, and tail as Nanosaurus? rex, which had been damaged by other collectors prior to description. [12] By 1977, he had concluded that Nanosaurus agilis was quite different from N. rex and the new skeleton, and coined Othnielia for the latter species. The paper (primarily concerning the transcontinental nature of Dryosaurus ) considered Laosaurus consors and L. gracilis synonyms of O. rex without elaboration, and considered L. celer an invalid nomen nudum . [13]

In 1990, Robert Bakker, Peter Galton, James Siegwarth, and James Filla described remains of a dinosaur they named Drinker nisti. The name is somewhat ironic; Drinker, named for renowned palaeontologist Edward Drinker Cope whose infamous "bone wars" with rival Othniel Charles Marsh produced many dinosaur fossils which are world-famous today, was described as a probable close relative of Othnielia, named for Marsh. The species name refers to the National Institute of Standards and Technology (NIST). Discovered by Siegwarth and Filla in upper Morrison Formation beds at Como Bluff, Wyoming, it was based on a partial subadult skeleton (listed as CPS 106 originally, then as Tate 4001 by Bakker 1996 [14] ) including partial jaws, vertebrae, and partial limbs. Several other specimens found in the same area were assigned to it, mostly consisting of vertebral and hindlimb remains, and teeth. [15] The holotype specimen's current location is unknown; according to Carpenter and Galton (2018), the previous two institutions reported to have had it did not ever curate the specimen, and the collection it was originally said to be in never existed at all. [14]

Life restoration Othnielosaurus.jpg
Life restoration

Several decades later, in his 2007 study of the teeth of Morrison ornithischians, Galton concluded that the holotype femur of Othniela rex is not diagnostic, and reassigned the BYU skeleton to Laosaurus consors, which is based on better material. As the genus Laosaurus is also based on nondiagnostic material, he gave the species L. consors its own genus, Othnielosaurus. As a result, in practical terms, what had been thought of as Othnielia is now known as Othnielosaurus consors. Regarding Nanosaurus agilis, Galton considered it a potentially valid basal ornithopod, and noted similarities to heterodontosaurids in the thigh bone. He tentatively assigned to it some teeth that had been referred to Drinker . [4]

Another decade later, in 2018, Galton, alongside Kenneth Carpenter, described a new ornithischian specimen. They found it very similar to the fragmentary holotype of Nanosaurus, but more clear in its anatomical features. Their new specimen was also found to display extreme similarity with the specimens of Othnielosaurus and Drinker. Due to the new data, they concluded that all three species, alongside Othnielia, represented the same animal, united under the name Nanosaurus agilis. This painted a new picture of a singular, very common small dinosaur known from a large amount of material. [14] This conclusion has been recognized by papers since, some of which incorporating the new, all-encompassing taxon into their phylogenetic analyses. [16] [17] [18]

Classification

The cladogram below results from analysis by Herne et al., 2019. [16]

Ornithischia

Heterodontosauridae

Eocursor

Thyreophora

Neornithischia

Lesothosaurus

Agilisaurus

Hexinlusaurus

Yandusaurus

Nanosaurus

Jeholosauridae

Haya

Jeholosaurus

Changchunsaurus

Thescelosauridae

Orodromeus

Koreanosaurus

Zephyrosaurus

Yueosaurus

Thescelosaurus

Cerapoda

Marginocephalia

Ornithopoda

Paleobiology and paleoecology

Casts mounted as if fighting, Wyoming Dinosaur Center Fighting Othnielia.jpg
Casts mounted as if fighting, Wyoming Dinosaur Center

Nanosaurus was one of the smaller members of the diverse Morrison Formation dinosaur fauna, diminutive in comparison to the giant sauropods. [19] The Morrison Formation is interpreted as a semiarid environment with distinct wet and dry seasons, and flat floodplains. [20] Vegetation varied from river-lining gallery forests of conifers, tree ferns, and ferns, to fern savannas with rare trees. [21] It has been a rich fossil hunting ground, holding fossils of green algae, fungi, mosses, horsetails, ferns, cycads, ginkgoes, and several families of conifers. Other fossils discovered include bivalves, snails, ray-finned fishes, frogs, salamanders, turtles, sphenodonts, lizards, terrestrial and aquatic crocodylomorphs, several species of pterosaur, numerous dinosaur species, and early mammals such as docodonts, multituberculates, symmetrodonts, and triconodonts. Such dinosaurs as the theropods Ceratosaurus , Allosaurus , Ornitholestes , and Torvosaurus , the sauropods Apatosaurus , Brachiosaurus , Camarasaurus , and Diplodocus , and the ornithischians Camptosaurus , Dryosaurus , and Stegosaurus are known from the Morrison. [22] Nanosaurus is present in stratigraphic zones 2-5. [23]

Typically, Nanosaurus has been interpreted like other hypsilophodonts as a small, swift herbivore, [24] although Bakker (1986) interpreted Nanosaurus as an omnivore. [25]

Related Research Articles

<i>Allosaurus</i> Extinct genus of carnosaurian theropod dinosaur

Allosaurus is an extinct genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic period. The name "Allosaurus" means "different lizard", alluding to its unique concave vertebrae. It is derived from the Greek words ἄλλος and σαῦρος. The first fossil remains that could definitively be ascribed to this genus were described in 1877 by famed paleontologist Othniel Charles Marsh. As one of the first well-known theropod dinosaurs, it has long attracted attention outside of paleontological circles.

<i>Stegosaurus</i> Thyreophoran stegosaurid dinosaur genus from Late Jurassic period

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 found in the western United States and in Portugal, where they are found in Kimmeridgian- to Tithonian-aged strata, dating to between 155 and 145 million years ago. Of the species that have been classified in the upper Morrison Formation of the western US, only three are universally recognized: S. stenops, S. ungulatus and S. sulcatus. The remains of over 80 individual animals of this genus have been found. Stegosaurus would have lived alongside dinosaurs such as Apatosaurus, Diplodocus, Camarasaurus and Allosaurus, the latter of which may have preyed on it.

<span class="mw-page-title-main">Othniel Charles Marsh</span> American paleontologist

Othniel Charles Marsh was an American professor of paleontology at Yale College and president of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among his legacies are the discovery or description of dozens of new species and theories on the origins of birds.

<i>Camarasaurus</i> Camarasaurid sauropod dinosaur genus from Late Jurassic Period

Camarasaurus was a genus of quadrupedal, herbivorous dinosaurs and is the most common North American sauropod fossil. Its fossil remains have been found in the Morrison Formation, dating to the Late Jurassic epoch, between 155 and 145 million years ago.

<i>Anchisaurus</i> Extinct genus of dinosaurs

Anchisaurus is a genus of basal sauropodomorph dinosaur. It lived during the Early Jurassic Period, and its fossils have been found in the red sandstone of the Portland Formation, Northeastern United States, which was deposited from the Hettangian age into the Sinemurian age, between about 200 and 195 million years ago. Until recently it was classed as a member of Prosauropoda. The genus name Anchisaurus comes from the Greek αγχιanchi-; "near, close" + Greek σαυρος ; "lizard". Anchisaurus was coined as a replacement name for "Amphisaurus", which was itself a replacement name for Hitchcock's "Megadactylus", both of which had already been used for other animals.

<span class="mw-page-title-main">Ornithopoda</span> Extinct suborder of dinosaurs

Ornithopoda is a clade of ornithischian dinosaurs, called ornithopods, that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world, dominating the North American land. Their major evolutionary advantage was the progressive development of a chewing apparatus that became the most sophisticated ever developed by a non-avian dinosaur, rivaling that of modern mammals such as the domestic cow. They reached their apex of diversity and ecological dominance in the hadrosaurids, before they were wiped out by the Cretaceous–Paleogene extinction event along with all other non-avian dinosaurs. Members are known from all seven continents, though they are generally rare in the Southern Hemisphere.

<i>Barosaurus</i> Diplodocid sauropod dinosaur genus from Upper Jurassic Period

Barosaurus was a giant, long-tailed, long-necked, plant-eating sauropod dinosaur closely related to the more familiar Diplodocus. Remains have been found in the Morrison Formation from the Upper Jurassic Period of Utah and South Dakota. It is present in stratigraphic zones 2–5.

<i>Dryosaurus</i> Extinct genus of dinosaurs

Dryosaurus is a genus of an ornithopod dinosaur that lived in the Late Jurassic period. It was an iguanodont. Fossils have been found in the western United States and were first discovered in the late 19th century. Valdosaurus canaliculatus and Dysalotosaurus lettowvorbecki were both formerly considered to represent species of Dryosaurus.

<i>Camptosaurus</i> Extinct genus of dinosaurs

Camptosaurus is a genus of plant-eating, beaked ornithischian dinosaurs of the Late Jurassic period of western North America and possibly also Europe. The name means 'flexible lizard'.

<span class="mw-page-title-main">Como Bluff</span> United States historic place

Como Bluff is a long ridge extending east–west, located between the towns of Rock River and Medicine Bow, Wyoming. The ridge is an anticline, formed as a result of compressional geological folding. Three geological formations, the Sundance, the Morrison, and the Cloverly Formations, containing fossil remains from the Late Jurassic of the Mesozoic Era are exposed.

<span class="mw-page-title-main">Hypsilophodontidae</span> Extinct family of dinosaurs

Hypsilophodontidae is a traditionally used family of ornithopod dinosaurs, generally considered invalid today. It historically included many small bodied bipedal neornithischian taxa from around the world, and spanning from the Middle Jurassic until the Late Cretaceous. This inclusive status was supported by some phylogenetic analyses from the 1990s and mid 2000s, although there have also been many finding that the family is an unnatural grouping which should only include the type genus, Hypsilophodon, with the other genera being within clades like Thescelosauridae and Elasmaria. A 2014 analysis by Norman recovered a grouping of Hypsilophodon, Rhabdodontidae and Tenontosaurus, which he referred to as Hypsilophodontia. All other analyses from around the same time have instead found these latter taxa to be within Iguanodontia.

Coelurus is a genus of coelurosaurian dinosaur from the Late Jurassic period. The name means "hollow tail", referring to its hollow tail vertebrae. Although its name is linked to one of the main divisions of theropods (Coelurosauria), it has historically been poorly understood, and sometimes confused with its better-known contemporary Ornitholestes. Like many dinosaurs studied in the early years of paleontology, it has had a confusing taxonomic history, with several species being named and later transferred to other genera or abandoned. Only one species is currently recognized as valid: the type species, C. fragilis, described by Othniel Charles Marsh in 1879. It is known from one partial skeleton found in the Morrison Formation of Wyoming, United States. It was a small bipedal carnivore with elongate legs.

<i>Alocodon</i> Extinct genus of dinosaurs

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<i>Laosaurus</i> Extinct genus of dinosaurs

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<i>Phyllodon</i> Extinct genus of dinosaurs

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<i>Camarasaurus grandis</i> Extinct species of dinosaur

Camarasaurus grandis is an extinct species of sauropod dinosaur in the genus that lived during the Jurassic in what is now the western United States. It is the geologically oldest of the four species of the genus Camarasaurus.

Eobatrachus is a dubious genus of extinct frog known only from the holotype, YPM 1862, part of the right humerus, found in Reed's Quarry 9 near Como Bluff, Wyoming in the Late Jurassic-aged Morrison Formation. The type, and only species, E. agilis, was named by Othniel Charles Marsh in 1887 and he initially interpreted it as a mammal, although it was later re-classified as a genus of frog related to Comobatrachus and Eobatrachus is now seen as a dubious amphibian genus, possibly belonging to Anura (frogs) according to Foster (2007).

<i>Uteodon</i> Genus of reptiles (fossil)

Uteodon is a genus of herbivorous iguanodontian dinosaur. It is a basal iguanodontian which lived during the late Jurassic period in what is now Uintah County, Utah. It is known from the middle of the Brushy Basin Member, Morrison Formation. The genus was named by Andrew T. McDonald in 2011 and the type species is U. aphanoecetes.

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