''Carusia'' is an extinct

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 ...

lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...

s from 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 ...

of Mongolia. It is a close relative of the family

Xenosauridae Xenosauridae is a family of anguimorph lizards whose only living representative is the genus '' Xenosaurus'', which is native to Central America. Xenosauridae also includes the extinct genera '' Exostinus'' and '' Restes''. Also known as knob-scal ...

, which includes living knob-scaled lizards. Fossils of the type and only species ''Carusia intermedia'' come from the 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. Campanian s ...

age

Barun Goyot Formation The Barun Goyot Formation (also known as Baruungoyot Formation or West Goyot Formation) 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 Pro ...

and have been found in the

Flaming Cliffs The Flaming Cliffs site (also known as Bayanzag ( zh, 巴彥扎格), Bain-Dzak or Bayn Dzak) ( mn, Баянзаг ''rich in saxaul''), with the alternative Mongolian name of mn, Улаан Эрэг (''red cliffs''), is a region of the Gobi Desert ...

, Ukhaa Tolgod, and Kheerman Tsav fossil localities. ''Carusia'' was first described in 1985 under the name ''Carolina intermedia'', but since the name ''Carolina'' was preoccupied by a genus of

scarab beetles The family Scarabaeidae, as currently defined, consists of over 30,000 species of beetles worldwide; they are often called scarabs or scarab beetles. The classification of this family has undergone significant change in recent years. Several sub ...

that had been named in 1880, it was renamed ''Carusia intermedia''. ''Carusia'' had initially been known from fragmentary skull material, complicating efforts to determine its evolutionary relationships with other lizards; it had variously been described as an indeterminate

scincomorph Scincomorpha is an infraorder and clade of lizards including skinks (Scincidae) and their close relatives. These include the living families Cordylidae (girdled lizards), Gerrhosauridae (plated lizards), and Xantusiidae (night lizards), as wel ...

, a xenosaurid, or some other type of

autarchoglossa Autarchoglossa is a clade (evolutionary grouping) of squamates that includes skinks, anguimorphs, snakes, and relatives. Autarchoglossa is supported as a monophyletic grouping (i.e. a valid clade) by morphological features in living and extinct l ...

n lizard convergent with xenosaurids. However, the discovery of 35 complete skulls in the 1990s, three of which were described in a detailed 1998 monograph, revealed that ''Carusia'' was the

sister taxon 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 t ...

(closest relative) of Xenosauridae, compelling the authors of the monograph to create a new clade called

Carusioidea Carusioidea is a clade of lizards that includes the family Xenosauridae (knob-scaled lizards) from Central America and the extinct genus '' Carusia'' from the Late Cretaceous of Mongolia. It was named in 1998 after a sister-group relationship wa ...

to include both taxa. Like xenosaurids, ''Carusia'' has a

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 ...

covered in large rounded

osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...

s (bony plates embedded in the skin). It also shares with xenosaurids closely spaced

orbits In celestial mechanics, an orbit 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 object or position in space such as a p ...

(eye sockets) with fused

frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...

s between them, and a connection between the

jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...

and

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 co ...

s. However, many other features of its skull set it apart from xenosaurids, including the lack of a

lacrimal bone The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...

, the wideness of the

palatine bone In anatomy, the palatine bones () are two irregular bones of the facial skeleton in many animal species, located above the uvula in the throat. Together with the maxillae, they comprise the hard palate. (''Palate'' is derived from the Latin ''pa ...

, and the small size and high number of teeth in its jaws.

References

Carusioidea Cretaceous lizards Lizard genera Late Cretaceous lepidosaurs of Asia Fossil taxa described in 1985 {{paleo-lizard-stub