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''Abrictosaurus'' (; "wakeful 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 heterodontosaurid
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 ...
that lived during the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic� ...
in what is now in parts of southern
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 ...
such as
Lesotho Lesotho, formally the Kingdom of Lesotho and formerly known as Basutoland, is a landlocked country in Southern Africa. Entirely surrounded by South Africa, it is the largest of only three sovereign enclave and exclave, enclaves in the world, t ...
and
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. It was a
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
herbivore 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 ...
or
omnivore 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 t ...
and was one of the most basal heterodontosaurids. It was approximately long and weighed less than . This dinosaur is known from the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
remains of only two individuals, found in the Upper Elliot Formation of Qacha's Nek District in Lesotho and
Cape Province The Province of the Cape of Good Hope (), commonly referred to as the Cape Province () and colloquially as The Cape (), was a province in the Union of South Africa and subsequently the Republic of South Africa. It encompassed the old Cape Co ...
in South Africa. The Upper Elliot is thought to date from the
Hettangian The Hettangian is the earliest age and lowest stage of the Jurassic Period of the geologic timescale. It spans the time between 201.3 ± 0.2 Ma and 199.3 ± 0.3 Ma (million years ago). The Hettangian follows the Rhaetian (part of the Triass ...
and
Sinemurian In the geologic timescale, the Sinemurian is an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series. It spans the time between 199.5 ±0.3 annu ...
stages of the Early Jurassic Period, approximately 200 to 190
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
. This formation is thought to preserve
sand dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
s as well as seasonal
floodplains A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high discharge.Goudie, ...
, in a semiarid environment with sporadic rainfall. Other dinosaurs found in this formation include the
theropod 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 ...
''
Megapnosaurus ''Megapnosaurus'' (meaning "big dead lizard", from Greek μέγα = "big", ἄπνοος = "not breathing", "dead", σαῦρος = "lizard") is an extinct genus of coelophysid theropod dinosaur that lived approximately 188 million years ago du ...
'', the
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
''
Massospondylus ''Massospondylus'' ( ) is a genus of sauropodomorph dinosaur from the Early Jurassic of southern Africa. It was described by Sir Richard Owen in 1854 from remains discovered in South Africa, and is thus one of the first dinosaurs to have been ...
'', as well as other heterodontosaurids like ''
Heterodontosaurus ''Heterodontosaurus'' is a genus of heterodontosaurid dinosaur that lived during the Early Jurassic, 200–190 million years ago. Its only known member species, ''Heterodontosaurus tucki'', was named in 1962 based on a skull discovered in South ...
'' and '' Lycorhinus''. Remains of terrestrial crocodylomorphs,
cynodont Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
s and early
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s are also abundant.


Description

Heterodontosaurids Heterodontosauridae is a family (biology), family of ornithischian dinosaurs that were likely among the most Basal (phylogenetics), basal (primitive) members of the group. Their phylogenetic placement is uncertain but they are most commonly fou ...
like ''Abrictosaurus'' were small, early
ornithischia Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
ns, named for their markedly
heterodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. Human dentition is heterodont and diphyodont as an example. In vertebrates, heterodont pertains to animals wher ...
dentition. They are best known for the large, canine-like tusks (often called ''
caniniform In mammalian oral anatomy, the canine teeth, also called cuspids, dogteeth, eye teeth, vampire teeth, or fangs, are the relatively long, pointed tooth, teeth. In the context of the upper jaw, they are also known as ''fangs''. They can appear mo ...
s'') in both upper and lower jaws. There were no teeth in the front of the jaws, where a hard
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for pecking, grasping, and holding (in probing for food, eating, manipulating and ...
was used to crop vegetation. There were three
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
ry teeth, with the first two small and conical and the third enlarged to form the upper caniniform, counterpart to the even larger lower caniniform, which was the first
dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
tooth. In the upper jaw, a large gap (or
diastema A diastema (: diastemata, from Greek , 'space') is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition may be referred to ...
) accommodated the lower caniniform tooth and separated the premaxillary teeth from the wider chewing teeth of the
maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
. Similar teeth lined the remainder of the lower jaw. ''Abrictosaurus'' is usually considered the most basal member of the family Heterodontosauridae. ''Lycorhinus'' and ''Heterodontosaurus'' both had high-crowned cheek teeth, which overlapped each other in the jaw, forming a continuous chewing surface
analogous Analogy is a comparison or correspondence between two things (or two groups of things) because of a third element that they are considered to share. In logic, it is an inference or an argument from one particular to another particular, as oppose ...
to those of
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 ...
hadrosaurid Hadrosaurids (), also hadrosaurs or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod fami ...
s. ''Abrictosaurus'' had more widely separated cheek teeth, with lower crowns, more similar to other early ornithischians. It has been suggested that ''Abrictosaurus'' lacked tusks and that this is another primitive feature. However, caniniforms were clearly present on one of the two specimens of ''Abrictosaurus''. The upper caniniform measured 10.5 millimeters (0.4 inches) high, while the lower reached 17 mm (0.67 in). These caniniforms were serrated only on the
anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
surface, unlike those of ''Lycorhinus'' and ''Heterodontosaurus'', which were serrated on both anterior and posterior edges. ''Abrictosaurus'' also had smaller, less powerful forelimbs than ''Heterodontosaurus'' and one fewer
phalanx bone The phalanges (: phalanx ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. Structu ...
in both the fourth and fifth digits of the forelimb.


History and naming

Both specimens of ''Abrictosaurus'' are housed in the collection of
University College London University College London (Trade name, branded as UCL) is a Public university, public research university in London, England. It is a Member institutions of the University of London, member institution of the Federal university, federal Uni ...
. The
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 ...
specimen was discovered in Lesotho and consists of a partial
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
and skeleton (UCL B54).
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 ...
Richard Thulborn, who first described the specimen in 1974, considered it a new
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), ...
of ''Lycorhinus'' and named it ''L. consors'', using the
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 ...
word ' which means 'companion' or 'spouse'. As UCL B54 lacked the caniniforms which had been found in the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
, ''Lycorhinus angustidens'', Thulborn believed it to be female. Neither the skull nor the skeletons of ''Abrictosaurus'' have been fully described in the literature. A tooth from the latest
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
of
Switzerland Switzerland, officially the Swiss Confederation, is a landlocked country located in west-central Europe. It is bordered by Italy to the south, France to the west, Germany to the north, and Austria and Liechtenstein to the east. Switzerland ...
has been assigned to ''Abrictosaurus'' sp., but this has not been supported, as the specimen does not have unique characteristics of ''Abrictosaurus'', heterodontosaurids, or ornithischians in general. In 1975,
James Hopson James Allen Hopson (born 1935) is an American paleontologist and professor (now retired) at the University of Chicago. His work has focused on the evolution of the synapsids (a group of amniotes that includes the mammals), and has been focused on ...
redescribed a fragmentary heterodontosaur skull (UCL A100) found in
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
that Thulborn had previously assigned to ''Lycorhinus angustidens''. After showing that UCL A100 could not belong to ''L. angustidens'' but was instead more similar to UCL B54, Hopson erected a new genus to contain both specimens. The generic name ''Abrictosaurus'' (from the
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 ...
' (') meaning 'wakeful' and (') meaning 'lizard') refers to Hopson's disagreement with Thulborn's
hypothesis A hypothesis (: hypotheses) is a proposed explanation for a phenomenon. A scientific hypothesis must be based on observations and make a testable and reproducible prediction about reality, in a process beginning with an educated guess o ...
that heterodontosaurids underwent periods of
aestivation Aestivation ( (summer); also spelled estivation in American English) is a state of animal dormancy, similar to hibernation, although taking place in the summer rather than the winter. Aestivation is characterized by inactivity and a lowered m ...
(
hibernation Hibernation is a state of minimal activity and metabolic reduction entered by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It is mos ...
during hot and/or dry seasons). The specific name was retained, creating the new
binomial Binomial may refer to: In mathematics *Binomial (polynomial), a polynomial with two terms *Binomial coefficient, numbers appearing in the expansions of powers of binomials *Binomial QMF, a perfect-reconstruction orthogonal wavelet decomposition * ...
''Abrictosaurus consors''. Despite Hopson's renaming, Thulborn continued to consider ''Lycorhinus angustidens'', ''Heterodontosaurus tucki'', and ''Abrictosaurus consors'' to be three species of the genus ''Lycorhinus''. Most paleontologists maintain all three genera separately, although there is no precise definition of a species or genus in paleontology.


Sexual dimorphism

The hypothesis of
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 ...
in heterodontosaurids has long centered on ''Abrictosaurus''. Tusks are a sexually dimorphic trait in many modern mammals, including
musk deer Musk deer can refer to any one, or all eight, of the species that make up ''Moschus'', the only extant genus of the family (biology), family Moschidae. Despite being commonly called deer, they are not true deer belonging to the family Cervidae, b ...
,
walrus The walrus (''Odobenus rosmarus'') is a large pinniped marine mammal with discontinuous distribution about the North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere. It is the only extant species in the family Odobeni ...
,
Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living ''Elephas'' species. It is the largest living land animal in Asia and the second largest living Elephantidae, elephantid in the world. It is char ...
s and many
pigs The pig (''Sus domesticus''), also called swine (: swine) or hog, is an omnivorous, domesticated, even-toed, hoofed mammal. It is named the domestic pig when distinguishing it from other members of the genus '' Sus''. Some authorities cons ...
, with tusks being present primarily in males. The lack of tusks in UCL B54 led to suggestions that it was female; perhaps even a female of another species. The discovery of caniniforms in UCL A100 showed that ''A. consors'' also has this 'male' characteristic, suggesting that it is at least a valid species in its own right. However, UCL B54 may actually be a juvenile, based on its short face and unfused sacral (hip)
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e. Therefore, the lack of tusks could be a juvenile trait instead of a
secondary sexual characteristic A secondary sex characteristic is a physical characteristic of an organism that is related to or derived from its sex, but not directly part of its reproductive system. In humans, these characteristics typically start to appear during puberty ...
, weakening the case for sexual dimorphism.


References

{{Taxonbar, from=Q131598, from2=Q18511066 Heterodontosauridae Dinosaur genera Sinemurian dinosaurs Elliot Formation Dinosaurs of Lesotho Fossil taxa described in 1975 Taxa named by James Hopson