''Goyocephale'' is an

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

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 ...

pachycephalosauria Pachycephalosauria (; from Greek παχυκεφαλόσαυρος for 'thick headed lizards') is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachycephal ...

ornithischia Ornithischia () is an extinct order 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 s ...

n that lived in

Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million, ...

during the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...

about 76 million years ago. It was first described in 1982 by

Altangerel Perle Altangerel Perle (born 1945) is a Mongolian palaeontologist. He is employed at the National University of Mongolia. He has described species such as ''Goyocephale lattimorei'', '' Achillobator giganticus'' and '' Erlikosaurus andrewsi''. He has ...

Teresa Maryańska Teresa Maryańska (1937 – 3 October 2019) was a Polish paleontologist who specialized in Mongolian dinosaurs, particularly pachycephalosaurians and ankylosaurians. Peter Dodson (1998 p. 9) states that in 1974 Maryanska together with Halszka ...

and

Halszka Osmólska Halszka Osmólska (September 15, 1930 – March 31, 2008) was a Polish paleontologist who had specialized in Mongolian dinosaurs. Biography She was born in 1930 in Poznań. In 1949, she began to study biology at Faculty of Biology and Earth Scie ...

for a disarticulated skeleton with most of a

skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...

, part of the

forelimb A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...

and

hindlimb A hindlimb or back limb is one of the paired articulated appendages (limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-Hindl ...

, some of the

pelvic girdle The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The p ...

, and some

vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...

. Perle ''et al.'' named the remains ''Goyocephale lattimorei'', from the Mongolian гоё (''goyo''), meaning "decorated", and the

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 ...

κεφαλή (''kephale''), for head. The

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 ...

name honours

Owen Lattimore Owen Lattimore (July 29, 1900 – May 31, 1989) was an American Orientalist and writer. He was an influential scholar of China and Central Asia, especially Mongolia. Although he never earned a college degree, in the 1930s he was editor of ''Pacif ...

Description

''Goyocephale'' is known from a partial

, including both

mandibles In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...

, the

skull roof The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...

, part of the

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 ...

, part of the

braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...

region, the posterior skull, the

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 mammal has b ...

, and the

maxilla The maxilla (plural: ''maxillae'' ) in vertebrates 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 t ...

. The posterior edge of the skull roof, at the edge of the

squamosal bone The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral c ...

s, has many small bony bumps, which would have been the base of small horns in life. A feature shared with pachycephalosaurids, ''Goyocephale'' had a heterodont dentition, with large

caniniform In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...

premaxilla teeth, followed by a

diastema A diastema (plural diastemata, from Greek διάστημα, 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 ...

between the premaxilla and the maxilla, and the regular sub-triangular teeth in the maxilla. Teeth in the premaxilla become larger farther posterior, with the last being the largest. The mandible teeth are similar, with the first tooth being caniniform, and the remainder low and sub-triangular. Of the vertebrae, only the

atlas bone In anatomy, the atlas (C1) is the most superior (first) cervical vertebra of the spine and is located in the neck. It is named for Atlas of Greek mythology because, just as Atlas supported the globe, it supports the entire head. The atlas is ...

(the first

cervical vertebra 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 sau ...

) and the

sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...

and

tail vertebrae The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordate ...

are preserved. The sacrum includes four vertebrae, which are not strongly fused and incomplete. On the

ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...

surface of the second sacral

centrum (Latin for ''center'') may refer to: Places In Greenland * Nuuk Centrum, a district of Nuuk, Greenland * Centrum Lake, Greenland In the Netherlands * Amsterdam-Centrum, the inner-most borough of Amsterdam, Netherlands * Rotterdam Centrum, a borou ...

is a length-wise ridge, with a groove along its midline. The first and second tail vertebrae were found articulated with the sacrum, although they are much weakly fused than the sacral vertebrae. In addition, they can be distinguished from the sacrum because they possess zygapophyses that are not fused together, although they also preserve a ventral groove. The limb and girdle bones are well-preserved, and show many features. The

humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...

is strongly bowed, with a nearly equal expansion on the

distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...

and proximal ends. It also shows a thick but weak deltopectoral crest (anterior projection on the humerus), and weakly separated

condyles A condyle (;Entry "condyle"
in
ilium bone shows a morphology typical for pachycephalosaurs, with a thin and horizontally elongate preacetabular process (front extension of the ilium), and a wide ridge extending outwards from the top edge of the postacetabular process (hind ilium extension). Also, the two preacetabular processes are strongly diverged in dorsal view, and in lateral view the ilium is almost straight and the postacetabular process is sub-rectangular. The

tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...

shows a pachycephalosaur design, and no

tarsal bones In the human body, the tarsus is a cluster of seven articulating bones in each foot situated between the lower end of the tibia and the fibula of the lower leg and the metatarsus. It is made up of the midfoot ( cuboid, medial, intermediate, and ...

appear to be articulated. The foot of ''Goyocephale'' is partially preserved, and at least three digits (digits II, III and IV) were present. On each toe, the

unguals An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropods and horned dinosaurs. A claw is a highly modified ungual ...

are triangular but not recurved, with the ungulate of the third toe being the largest.

Classification

''Goyocephale'' is a primitive

n, and was originally included in the family Homalocephalidae, which united the genus with ''

Homalocephale ''Homalocephale'' (from Ancient Greek, Greek ὁμαλός, ''homalos'', "even", and κεφαλή, ''kephalē'', "head") is a genus of pachycephalosaurid dinosaur that lived during the Late Cretaceous period of what is now the Nemegt Formation, M ...

'', which also has a flat skull roof. ''Goyocephale'' is distinguished from ''Homalocephale'' by its overall proportions, the shape of its supratemporal fenestra, and heterodont dentition, although the two share multiple features. However, many more recent

phylogenetic analyses In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

tend to find Homalocephalidae to be a

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

family, with the genera included simply being consecutive branches sister to Pachycephalosauridae, or as consecutive branches primitive to ''

Prenocephale ''Prenocephale'' (meaning "sloping head") is a genus of small pachycephalosaurid dinosaur from the Late Cretaceous Nemegt Formation of Mongolia. It was similar in many ways to its close relative, ''Homalocephale''. Description Adult ''Prenoceph ...

'' but within Pachycephalosauridae. A

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

illustrating the latter hypothesis is shown below. Goyocephalebcn2

However, one phylogenetic analysis did support a

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

Homalocephalidae, with ''Goyocephale'', ''Homalocephale'' and ''Wannanosaurus'' being the most derived pachycephalosaurians, and their sister taxa being ''Prenocephale'' and ''

Tylocephale ''Tylocephale'' (meaning "swollen head", from the Greek ''τυλη'' meaning 'callus' or 'hard swelling' and ''κεφαλη'' meaning 'head') is a genus of pachycephalosaurid dinosaur from the Late Cretaceous period. It was a herbivorous dinosau ...

''. This positioning of all the taxa from Asia as most derived would support that pachycephalosaurians evolved in North America, where they retained their greatest diversity, before migrating to Asia. In addition, the oldest genera that belong in the group are North American, which provides further support for this position.

References

{{Taxonbar, from=Q133079 Late Cretaceous dinosaurs of Asia Pachycephalosaurs Fossil taxa described in 1982 Taxa named by Altangerel Perle Taxa named by Halszka Osmólska Taxa named by Teresa Maryańska Ornithischian genera

Description

Classification

See also

References