Dysalotosaurus
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''Dysalotosaurus'' ("uncatchable 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
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 ...
iguanodont Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivore, herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and rel ...
ian
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 ...
. It was a
dryosaurid Dryosauridae was a family of primitive iguanodonts, first proposed by Milner & Norman in 1984. They are known from Middle Jurassic to Early Cretaceous rocks of Africa, Europe, and North America. Dryosauridae was first proposed in 1984 by Britis ...
iguanodontian, and its fossils have been found in late
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 154.8 ±0.8 Ma and 149.2 ±0.7 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
-age rocks (
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 ...
) of the
Tendaguru Formation The Tendaguru Formation, or Tendaguru Beds are a highly List of fossiliferous stratigraphic units in Tanzania, fossiliferous Lithostratigraphy, formation and Lagerstätte located in the Lindi Region of southeastern Tanzania. The formation represe ...
of
Lindi Region Lindi Region (''Mkoa wa Lindi'' in Swahili language, Swahili) is one of Tanzania's 31 administrative Regions of Tanzania, regions. The region covers an area of . The region is comparable in size to the combined land area of the nation state of Sr ...
in
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
. 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 species of the genus is ''D. lettowvorbecki''. This species was named by
Hans Virchow Hans may refer to: __NOTOC__ People * Hans (name), a masculine given name * Hans Raj Hans, Indian singer and politician ** Navraj Hans, Indian singer, actor, entrepreneur, cricket player and performer, son of Hans Raj Hans ** Yuvraj Hans, Punjabi a ...
in
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 ...
in honor of the
Imperial German (, literally translated ) is an archaic term for those ethnic Germans who resided within the German state that was founded in 1871. In contemporary usage, it referred to German citizens, the word signifying people from the German ', i.e., Imp ...
Army Officer,
Paul von Lettow-Vorbeck Paul Emil von Lettow-Vorbeck (20 March 1870 – 9 March 1964), popularly known as the Lion of Africa (), was a general in the Imperial German Army and the commander of its forces in the German East Africa campaign. For four years, with a force ...
. For much of the 20th century the species was referred to the related and approximately contemporary genus ''
Dryosaurus ''Dryosaurus'' ( , meaning 'tree lizard', Greek ' () meaning 'tree, oak' and () meaning 'lizard', (the name reflects the forested habitat, not a vague oak-leaf shape of its cheek teeth as is sometimes assumed) is a genus of an ornithopod dinos ...
'', but newer studies reject this synonymy.


Discovery and naming

Thousands of ''Dysalotosaurus'' bones and bone fragments have been recovered from the
Tendaguru Formation The Tendaguru Formation, or Tendaguru Beds are a highly List of fossiliferous stratigraphic units in Tanzania, fossiliferous Lithostratigraphy, formation and Lagerstätte located in the Lindi Region of southeastern Tanzania. The formation represe ...
in
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
,
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
since 1909; the specimens described by
Hans Virchow Hans may refer to: __NOTOC__ People * Hans (name), a masculine given name * Hans Raj Hans, Indian singer and politician ** Navraj Hans, Indian singer, actor, entrepreneur, cricket player and performer, son of Hans Raj Hans ** Yuvraj Hans, Punjabi a ...
(1919) were discovered between 1909 and 1913. It has been suggested that all of these specimens were part of a herd that was killed in a mass death event. The genus was named by Hans Virchow in 1919,Virchow, Hans (1919). Atlas und Epistropheus bei den Schildkröten. ''Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin'' 8: 303–332. as opposed to
Josef Felix Pompeckj Josef Felix Pompeckj (May 10, 1867, Groß-Köllen – July 8, 1930, Berlin) was a German paleontologist and geologist. He was born in Groß-Köllen, Kingdom of Prussia (now Kolno in Poland). He studied geology and paleontology at the University ...
(1920), who is often incorrectly cited as naming the genus.


Description

''Dysalotosaurus'' was a small, relatively basal iguanodontian
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
. It lacked the large thumb spikes found in later iguanodontians, and was more adapted for
bipedalism Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) Limb (anatomy), limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from ...
than its larger relatives with its short front limbs and long, counter-balancing tail. ''Dysalotosaurus'' had powerful and long hind limbs, suggesting it was relatively
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often ...
compared to ''
Iguanodon ''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species found worldwide have been classified in the genus ''Iguanodon'', dating from the Late Jurassic to Early Cretaceous, Taxonomy (bi ...
'' and other members of the clade. A recent comparative study based on bone cross-sectional geometry confirmed its bipedal locomotion with erect stance. Based on CT scans of the braincase, it is believed that ''Dysalotosaurus'' held its head dorsally (pointing straight forward) when not feeding. The same study also suggested, based on the morphology of the inner ear, that ''Dysalotosaurus'' was not able to discern between high- and low-frequency sounds (like most herbivorous dinosaurs). However, it also had other adaptations that are commonly associated with derived hearing abilities, rendering its sensory capabilities unclear. In 2016,
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
estimated ''Dysalotosaurus''' length at , and its weight at .


Ontogeny

Various specimens of ''Dysalotosaurus'' are known, all representing animals that were not yet
sexually mature Sexual maturity is the capability of an organism to reproduce. In humans, it is related to both puberty and adulthood. ''Puberty'' is the biological process of sexual maturation, while ''adulthood'', the condition of being socially recognized as ...
at the time of their deaths. Ontogenetic studies demonstrate typical aging trends, such as a lengthening of the snout and relative shrinking of the
orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
. Differences in
dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...
as the animals aged also suggest a change from an
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
diet early in life to fully
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 ...
feeding habits as an adult. This switch reflects the general evolutionary trend towards obligate herbivory among iguanodontians and other ornithopods. ''Dysalotosaurus'' also appears to have had a life expectancy of roughly twenty years and lived in herds of mixed ages.


Palaeobiology

''Dysalotosaurus'' was a precocial dinosaur, which experienced sexual maturity at ten years, had an indeterminate growth pattern, and maximum growth rates comparable to a large kangaroo. A tibia (SMNS T3) and a fibula (GPIT/RE/5109) of ''Dysalotosaurus'' preserve evidence of medullary bone tissue, which are found in modern birds that are laying eggs in reproduction, and in some other non-avian dinosaurs such as ''
Tyrannosaurus rex ''Tyrannosaurus'' () is a genus of large theropoda, theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It live ...
''.


Palaeopathology

In 2011 paleontologists Florian Witzmann and Oliver Hampe from the
Museum für Naturkunde The Natural History Museum () is a natural history museum located in Berlin, Germany. It exhibits a vast range of specimens from various segments of natural history and in such domain it is one of three major museums in Germany alongside Naturm ...
and colleagues discovered that deformations of some ''Dysalotosaurus'' bones were likely caused by a viral infection similar to
Paget's disease of bone Paget's disease of bone (commonly known as Paget's disease or, historically, osteitis deformans) is a condition involving Bone remodeling, cellular remodeling and deformity of one or more bones. The affected bones show signs of dysregulated bone ...
. This is the oldest evidence of viral infection known to science. The discovery of a hemivertebra (spinal malformation) in another specimen likely caused
scoliosis Scoliosis (: scolioses) is a condition in which a person's Vertebral column, spine has an irregular curve in the coronal plane. The curve is usually S- or C-shaped over three dimensions. In some, the degree of curve is stable, while in others ...
and other pathologic effects on the animal during its life. This malformation occurs in almost every vertebrate clade, though this is the first known evidence of the condition in dinosaurs.


Palaeoecology

Contemporary dinosaurs were ''
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 ...
'', various
sauropods Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
including ''
Giraffatitan ''Giraffatitan'' (name meaning "titanic giraffe") is a genus of sauropod dinosaur that lived during the late Jurassic Period (geology), Period (Kimmeridgian–Tithonian stages) in what is now Lindi Region, Tanzania. Only one species is known, ...
'' and ''
Dicraeosaurus ''Dicraeosaurus'' (Gr. , ' "bifurcated, double-headed" + Gr. , ' "lizard") is a genus of diplodocoid sauropod dinosaur that lived in what is now Lindi Region, Tanzania during the late Jurassic period. The genus was named for the neural spines on ...
'', and large
theropods Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
including ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Ancient Greek, Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic Epoch (Bathonian stage, 166 ...
'' and ''
Ceratosaurus ''Ceratosaurus'' (from Greek 'horn' and 'lizard') is a genus of carnivorous theropod dinosaur that lived in the Late Jurassic period (Kimmeridgian to Tithonian ages). The genus was first described in 1884 by American paleontologist Othni ...
''.
Pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s are also common in the Tendaguru Formation, as well as
mammaliaformes Mammaliaformes ("mammalian forms") is a clade of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined as ...
(ancestral to mammals). During the Jurassic, Tendaguru was part of the
semi-arid A semi-arid climate, semi-desert climate, or steppe climate is a aridity, dry climate sub-type. It is located on regions that receive precipitation below Evapotranspiration#Potential evapotranspiration, potential evapotranspiration, but not as l ...
coastline of
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 ...
. ''Dysalotosaurus'' fossils were discovered in
strata In geology and related fields, a stratum (: strata) is a layer of Rock (geology), rock or sediment characterized by certain Lithology, lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by v ...
that were deposited in structures characteristic of
tidal flats Mudflats or mud flats, also known as tidal flats or, in Ireland, slob or slobs, are coastal wetlands that form in intertidal areas where sediments have been deposited by tides or rivers. A global analysis published in 2019 suggested that tidal ...
and
lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') an ...
s.


References

{{Taxonbar, from=Q136892 Dryosauridae Dinosaur genera Kimmeridgian dinosaurs Tendaguru Formation Fossil taxa described in 1919 Dinosaurs of Tanzania