Zaraapelta Nomadis
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''Zaraapelta'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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
ankylosaurid Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
thyreophoran 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 ...
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 ...
from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
of
Mongolia Mongolia is a landlocked country in East Asia, bordered by Russia to the north and China to the south and southeast. It covers an area of , with a population of 3.5 million, making it the world's List of countries and dependencies by po ...
. 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 ...
is ''Zaraapelta nomadis'', named and described by Arbour ''et al'' in 2014. ''Zaraapelta'' is known from a single skull from the
Barun Goyot Formation The Baruungoyot Formation (also known as Barun Goyot) is a geological formation dating to the Late Cretaceous Period. It is located within and is widely represented in the Gobi Desert Basin, in the Ömnögovi Province of Mongolia. Descriptio ...
. It was found to be closest to ''
Tarchia ''Tarchia'' (meaning "brainy one") is a genus of herbivorous ankylosauridae, ankylosaurid dinosaur from the late Cretaceous of Mongolia. Discovery and naming In 1970, a Polish-Mongolian expedition discovered an ankylosaurian skull near Khulsan. ...
'' in the
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 ...
within its description.


Discovery and naming

In 2000, Robert Gabbard, member of a team headed by
Philip John Currie Philip John Currie (born March 13, 1949) is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the ...
, found an ankylosaur skull in the
Gobi Desert The Gobi Desert (, , ; ) is a large, cold desert and grassland region in North China and southern Mongolia. It is the sixth-largest desert in the world. The name of the desert comes from the Mongolian word ''gobi'', used to refer to all of th ...
near Hermiin Tsav at the Baruungoyot. In 2014, Victoria Megan Arbour named and described the find as the species ''Zaraapelta nomadis'', but at first it remained an invalid ''nomen ex dissertatione''.Arbour, Victoria Megan, 2014, ''Systematics, evolution, and biogeography of the ankylosaurid dinosaurs'' Ph.D thesis, University of Alberta Later that year, however, it was validly named as the type species ''Zaraapelta nomadis'' by Arbour, Currie and, posthumously, the female Mongolian paleontologist Demchig Badamgarav. The generic name is derived from
Mongolian Mongolian may refer to: * Something of, from, or related to Mongolia, a country in Asia * Mongolian people, or Mongols * Bogd Khanate of Mongolia, the government of Mongolia, 1911–1919 and 1921–1924 * Mongolian language * Mongolian alphabet * ...
', "hedgehog", in reference to the prickly appearance of ankylosaurs, and the Greek , ', "small shield", a common component of ankylosaurian names in view of their
body armour Body armour, personal armour (also spelled ''armor''), armoured suit (''armored'') or coat of armour, among others, is armour for human body, a person's body: protective clothing or close-fitting hands-free shields designed to absorb or deflect ...
. 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 the
genitive In grammar, the genitive case ( abbreviated ) is the grammatical case that marks a word, usually a noun, as modifying another word, also usually a noun—thus indicating an attributive relationship of one noun to the other noun. A genitive can ...
of 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 ...
', "nomad" and refers to the ''Nomadic Expeditions''
travel agency A travel agency is a private retailer or public service that provides travel and tourism-related services to the general public on behalf of accommodation or travel suppliers to offer different kinds of travelling packages for each destina ...
that has organised many palaeontological expeditions to Mongolia. 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 ...
, MPC D-100/1388, was found in a layer of the Barun Goyot Formation, dating from the mid to late
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
, about seventy-five million years old. It consists of a skull lacking the snout tip. No elements of the lower jaws were discovered. The specimen probably represents a subadult individual.


Description

The preserved skull has, lacking the
praemaxilla 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 has ...
e, about a length of forty centimetres, indicating that ''Zaraapelta'' was a medium-sized ankylosaurian. ''Zaraapelta'' differs from all other
Ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the clade Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs ...
in the possession of a squamosal horn with a very smooth bone texture along the upper cutting edge, abruptly changing into a rough surface at the inner and the outer side; and in the presence of an intricate pattern of a large number of osteoderms behind the eye socket. Beside these
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 ...
, ''Zaraapelta'' differs from its close relative ''
Saichania ''Saichania'' (Mongolian meaning "beautiful one") is a genus of herbivorous ankylosaurid dinosaur from the Late Cretaceous period of Mongolia. The first fossils of ''Saichania'' were found in the early 1970s in Mongolia. In 1977 the type speci ...
'' in having a large number of osteoderms in front of the eye socket; by a notch in the rim above the eye socket, causing the two supraorbital osteoderms there to have separate peaks; by the lack of distinct ', head armour tiles, behind the rear of the middle supraorbital; by less protruding osteoderms on the rear edge of the skull roof; and by the rear of the skull and the
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the Atlas (anatomy), atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape ...
being visible in top view. ''Zaraapelta'' differs from the relative ''
Tarchia ''Tarchia'' (meaning "brainy one") is a genus of herbivorous ankylosauridae, ankylosaurid dinosaur from the late Cretaceous of Mongolia. Discovery and naming In 1970, a Polish-Mongolian expedition discovered an ankylosaurian skull near Khulsan. ...
'' in having a less sideways protruding ' on the
prefrontal Prefrontal may refer to: *Prefrontal bone, a skull bone in some tetrapods *Prefrontal cortex, a region of the brain of a mammal *Prefrontal scales The prefrontal scales on snakes and other reptiles are the scales adjacent and anterior to the fr ...
; in the presence of a(n ossified) scroll-shaped turbinate bone in the nasal cavity, at the underside of the
frontal bone In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
; by the lack of distinct ', head armour tiles, behind the rear of the middle supraorbital; and by the fusion of the quadrate with the paroccipital process. The main difference with ''Tarchia'' resided in the unique squamosal horn shape. However, ''Tarchia'' itself has also a unique squamosal horn configuration in that an accessory osteoderm is present in front of it. It might thus in principle be possible that ''Zaraapelta'' simply represents an old ''Tarchia'' individual in which this osteoderm has shifted on top of the squamosal horn, creating the strange double-layered structure. This possibility was rejected by the authors however, because from other ankylosaurian species no comparably large
ontogenetic 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 egg to adult. The term can also be used to refer to the stu ...
changes of the squamosal horn shape are known. The squamosal horn, on the top rear corner of the skull, is robust and pyramid-shaped. There is a sharp cutting edge on top of the horn, bordered by smooth strips; these abruptly change into deeper rugose areas, so that rims are present on both sides. The quadratojugal cheeks horn is large with a concave rear edge. The cheek area between these large horns, behind the eye socket, is filled by flat osteoderms, separated by deep grooves, creating a delicate "dried mud" pattern. The eye socket itself is surrounded by a narrow bone ring. On its rear edge six osteoderms are present, those above rectangular in shape, the lower ones gradually becoming smaller and more triangular. Behind these are seven smaller tiles, roughly square-shaped, in an irregular pattern. The rear of this configuration is formed by a large triangular osteoderm. Just below the squamosal horn and above the cheek horn a row of small bumps is present.


Phylogeny

''Zaraapelta'' was placed in the
Ankylosauridae Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
. A
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analysis recovered it as the
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of ''Tarchia''. A phylogenetic analysis by Zheng ''et al''. (2018) led to the same result. Phylogenetic analyses by Xing ''et al''. (2024) recover it in slightly different positions within
Ankylosaurinae Ankylosaurinae is a subfamily of ankylosaurid dinosaurs, existing from the Early Cretaceous about 105 million years ago until the end of the Late Cretaceous, about 66 mya. Many genera are included in the clade, such as '' Ankylosaurus'', ''Pinac ...
. Below are two simplified
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 ...
s from that study: Topology A: Zheng ''et al''. (2018) dataset + ''
Datai ''Datai'' is an extinct genus of ankylosaurid dinosaur from the Late Cretaceous Zhoutian Formation of Jiangxi Province, China. The genus contains a single species, ''D. yingliangis'', which is known from the articulated skulls and partial skele ...
'' (14-taxon deletion) Topology B: Raven ''et al''. (2023) dataset + ''
Datai ''Datai'' is an extinct genus of ankylosaurid dinosaur from the Late Cretaceous Zhoutian Formation of Jiangxi Province, China. The genus contains a single species, ''D. yingliangis'', which is known from the articulated skulls and partial skele ...
'' (34-taxon deletion)


Paleobiology

Arbour pointed out that ''Saichania'', ''Tarchia'' and ''Zaraapelta'' shared the same habitat. She assumed that this relative ankylosaurian abundance in species had been caused by their being the main herbivores in the area, enough crop thus being available to feed three populations, although their relative niches were unclear. The differences in head ornamentation would then have served
species recognition Intra-species recognition is the recognition by a member of an animal species of a conspecific (another member of the same species). In many species, such recognition is necessary for procreation. Different species may employ different methods, ...
. She saw no indications 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 ...
. However, she also assumed that
sexual selection Sexual selection is a mechanism of evolution in which members of one sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex ...
had been a factor in the evolution of the typical ''Zaraapelta'' head ornamentation.


See also

*
Timeline of ankylosaur research This timeline of ankylosaur research is a chronological listing of events in the History of paleontology, history of paleontology focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spik ...


References

{{Taxonbar, from=Q18435455 Ankylosaurinae Dinosaur genera Maastrichtian dinosaurs Baruungoyot Formation Dinosaurs of Mongolia Fossil taxa described in 2014 Taxa named by Philip J. Currie