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''Rugops'' (meaning ‘wrinkle face’) is a
monospecific In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
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 ...
of basal
abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found ...
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
from
Niger ) , official_languages = , languages_type = National languagesLate 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 ...
period (
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
stage, ~95 Ma) in what is now the
Echkar Formation The Echkar Formation is a geological formation comprising sandstones and claystones in the Agadez Region of Niger, central Africa. Description Its strata date back to the Late Albian to Late Cretaceous ( Cenomanian stages, about 100-95 milli ...
. The type and only species, ''Rugops primus'', is known only from a partial
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 ...
. It was named and described in
2004 2004 was designated as an International Year of Rice by the United Nations, and the International Year to Commemorate the Struggle Against Slavery and its Abolition (by UNESCO). Events January * January 3 – Flash Airlines Flight 6 ...
by
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at sites ...
, Jeffery Wilson and Jack Conrad. ''Rugops'' has an estimated length of 4.4–5.3 metres (14.4–17.4 ft) and weight of 410 kilograms (900 lbs). The top of its skull bears several pits which correlates with overlaying scale and the front of the snout would have had an armour-like dermis.


Discovery and naming

A skull pertaining to an
abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found ...
was recovered during an expedition in 2000 led by
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at sites ...
near In-Abangharit, Niger Republic. The specimen came from the
Echkar Formation The Echkar Formation is a geological formation comprising sandstones and claystones in the Agadez Region of Niger, central Africa. Description Its strata date back to the Late Albian to Late Cretaceous ( Cenomanian stages, about 100-95 milli ...
of the Tegama Group which dates to the
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
stage of 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 ...
period, 96 Ma. The formation has also yielded specimens pertaining to the
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
'' Carcharodontosaurus iguidensis'',Brusatte, S.L. and Sereno, P.C. (2007). "A new species of ''Carcharodontosaurus'' (dinosauria: theropoda) from the Cenomanian of Niger and a revision of the genus." ''Journal of Vertebrate Paleontology'', 27(4): . the
spinosaurid The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
''
Spinosaurus ''Spinosaurus'' (; ) is a genus of spinosaurid dinosaur that lived in what now is North Africa during the Cenomanian to upper Turonian stages of the Late Cretaceous period, about 99 to 93.5 million years ago. The genus was known first fro ...
'', an unnamed rebbachisaurid and
titanosaur Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still th ...
, and the
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...
''
Fortignathus ''Fortignathus'' is an extinct genus of dyrosaurid or peirosaurid crocodylomorph Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to ...
'', ''
Laganosuchus ''Laganosuchus'' is an extinct genus of stomatosuchid crocodyliform. Fossils have been found from Niger and Morocco and date back to the Upper Cretaceous. Discovery The name means "pancake crocodile" from the Greek , ("pancake") and , ("cro ...
'' and ''
Kaprosuchus ''Kaprosuchus'' is an extinct genus of mahajangasuchid crocodyliform. It is known from a single nearly complete skull collected from the Upper Cretaceous Echkar Formation of Niger. The name means "boar crocodile" from the Greek , ''kapros'' ("bo ...
''. The specimen was described in 2004 by Paul Sereno, Jeffery Wilson and Jack Conrad. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
specimen, MNN IGU1, consists of a partial skull which lacks portions of the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
and
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 ...
. The type specimen may represent a subadult individual based on its small size, the lack of fusion between the
nasals In phonetics, a nasal, also called a nasal occlusive or nasal stop in contrast with an oral stop or nasalized consonant, is an occlusive consonant produced with a lowered velum, allowing air to escape freely through the nose. The vast majorit ...
and the presence of the
fenestra A fenestra (fenestration; plural fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomical st ...
between the prefrontal,
frontal Front may refer to: Arts, entertainment, and media Films * ''The Front'' (1943 film), a 1943 Soviet drama film * ''The Front'', 1976 film Music * The Front (band), an American rock band signed to Columbia Records and active in the 1980s and e ...
,
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
and
lacrimal bones 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 o ...
. The
generic Generic or generics may refer to: In business * Generic term, a common name used for a range or class of similar things not protected by trademark * Generic brand, a brand for a product that does not have an associated brand or trademark, other ...
name, ''Rugops'', is derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word "''ruga''" (wrinkle) and the
Greek Greek may refer to: Greece 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 ...
word "''opsi''" (face). The
specific Specific may refer to: * Specificity (disambiguation) * Specific, a cure or therapy for a specific illness Law * Specific deterrence, focussed on an individual * Specific finding, intermediate verdict used by a jury in determining the fina ...
name is derived from the Latin word "''primus''" (first). Both the generic and specific name refer to ''Rugops'' as being one of the earliest abelisaurids with a textured skull. In 2005, a partial right
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 ...
of an abelisaurid was described from the
Kem Kem Group The Kem Kem Group (commonly known as the Kem Kem beds) is a geological group in the Kem Kem region of eastern Morocco, whose strata date back to the Cenomanian stage of the Late Cretaceous. Its strata are subdivided into two geological formations ...
of
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
. The maxilla shares some similarities with ''Rugops'' such as with the morphology of the teeth, the shape of the alveolar, the rugose texture on the lateral sides, the straight border with 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 position of the palatal shelf. However, the maxilla cannot be referred to ''Rugops'' as the diagnostic features of the genus are located on different parts of the skull.


Description

In 2010,
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
gave ''Rugops'' an estimated length of 6 metres (19.7 ft) and weight of 750 kilograms (1,650 lbs). However, Grillo & Delcourt (2016) gave a lower estimate of 4.4 metres (14.4 ft) long while Molina-Pérez & Larramendi (2016) gave an estimate of 5.3 metres (17.4 ft) long and 410 kilograms (900 lbs) in weight.


Skull

The skull length of ''Rugops'' was about 31.5 centimetres (12.4 inches). As in other abelisaurids, the skull of ''Rugops'' has an external surface that is textured, a dental arcade that is U-shaped, maxillary-jugal contact that is broad, the presence of a socket on 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 ...
and
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * ...
that are subrectangular in shape. The orbital brow is present on the skull, although it is not fully formed. Many of the bones that make up the skull have slender proportions. In addition, the skull also has relatively thin
nasals In phonetics, a nasal, also called a nasal occlusive or nasal stop in contrast with an oral stop or nasalized consonant, is an occlusive consonant produced with a lowered velum, allowing air to escape freely through the nose. The vast majorit ...
and skull fenestrae the are proportionally large. The upper surface of each nasal bone has a row of several depressions with grooves for vascular supply going into them. A similar condition is seen in ''
Carnotaurus ''Carnotaurus'' (; ) is a genus of theropod dinosaur that lived in South America during the Late Cretaceous period, probably sometime between 71 and 69 million years ago. The only species is ''Carnotaurus sastrei''. Known from a single well-p ...
'', although the upper surface of the nasals is convex unlike ''Rugops'', as it is concave. Sereno ''et al.'' (2004) suggested that the depressions on the dorsal surface of each nasal anchored either sensory structures or soft tissues for display. Delcourt (2018), however, interpreted that they correlate with overlying
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
, as seen in extant
reptiles Reptiles, as most commonly defined are the animals in the Class (biology), class Reptilia ( ), a paraphyletic grouping comprising all sauropsid, sauropsids except birds. Living reptiles comprise turtles, crocodilians, Squamata, squamates (lizar ...
such as
crocodiles Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant memb ...
and
lizards 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 althou ...
. The anterior-most snout has a papillate texture which indicates the presence of an armour-like
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...
. The author suggested that, based on the type specimen probably being a subadult individual, the armour-like dermis may have reached a larger surface as it grew to which it developed a more papillate texture. Delcourt (2018) also proposed that the armour-like dermis may corelate with a low-motion headbutting behaviour, as seen in marine iguanas. A maxilla (NPSJB-PV247) from
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and gl ...
shows similarities with ''Rugops'' as the pattern of external ornamentation is nearly identical and the internal details are also have a close similarity. In ''Rugops'', the interdenticular sulci of the denticles, the presence of which might possible be a synapomorphic characteristic of Abelisauridae, is absent and has a higher dental formulae than in any other abelisaurid. The nasal sculpturing of ''Rugops'' is similar to the ornamentation seen in ''
Skorpiovenator ''Skorpiovenator'' ("scorpion hunter") is a genus of abelisaurid theropod dinosaur from the Late Cretaceous (Cenomanian to Turonian) Huincul Formation of Argentina. It is one of the most complete and informative abelisaurids yet known, describe ...
'' as the surface of the nasals show hummocky-like rugosities and has grooves that lead into each
foramen In anatomy and osteology, a foramen (;Entry "foramen"
in
. However, unlike ''Skorpiovenator'', ''Rugops'' lacks extra foramina on the skull roof that could represent homologues. The external morphology of the nasals are similar to that of ''Skorpiovenator'' as they share a similar foramina pattern. The ventral surface of the nasals have a series of foramina, which has been suggested to connect to an internal system. The describing authors indicated two distinguishing traits. Both of these are autapomorphies, unique derived characters. The skull roof has small fenestra that are present between the prefrontal, frontal, post-orbital and lacrimal. The dorsal surface of each nasal has a row of seven small depressions.


Classification

Sereno ''et al.'' (2004) initially found ''Rugops'' to be the basalmost abelisaurid, a position also recovered by various analyses by Egli ''et al.'' (2016), Delcourt (2018), Cerroni ''et al.'' (2020) and Rolando ''et al.'' (2020). However, ''Rugops'' has also been recovered as being more derived than ''
Rahiolisaurus'' and/or ''
Eoabelisaurus ''Eoabelisaurus'' () is a genus of abelisaurid theropod dinosaur from the Lower Jurassic Cañadón Asfalto Formation of the Cañadón Asfalto Basin in Argentina, South America. The generic name combines a Greek ἠώς, (''eos''), "dawn", with th ...
'' but more basal than other abelisaurids by Pol & Rauhut (2012), Rauhut & Carrano (2016), and Iori ''et al.'' (2021). Other alternative positions include ''Rugops'' being more derived than ''
Kryptops ''Kryptops'' is a genus of abelisaurid theropod dinosaur from the Early Cretaceous of Niger. It is known from a partial skeleton found at the Gadoufaoua locality in the western Ténéré Desert, in rocks of the Aptian-Albian age Elrhaz Formation. ...
'', ''
Chenanisaurus ''Chenanisaurus'' is a genus of predatory abelisaurid dinosaur, with a single known species ''C. barbaricus''. It comes from the upper Maastrichtian phosphates of the Ouled Abdoun Basin in Morocco, North Africa. The animal is known from a holo ...
'' and/or ''
Spectrovenator ''Spectrovenator'' ("ghost hunter", named because the holotype was unexpectedly found under the holotype of '' Tapuiasaurus'') is a genus of basal abelisaurid theropod dinosaur that lived during the early Cretaceous period in what is now Brazil. ...
'' as recovered by Sereno & Brusatte (2008), Zaher et al. (2020), Gianechini ''et al.'' (2021) and Agnolín ''et al.'' (2022), and within a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with other abelisaurids such as ''
Xenotarsosaurus ''Xenotarsosaurus'' is a genus of abelisaurid theropod dinosaur that lived during the Late Cretaceous of Argentina. Discovery In 1980 geologist Juan Carlos Sciutto discovered a rich fossil site six kilometres north of the Ocho Hermanos ranch in ...
'', '' Tarascosaurus'', '' Ilokelesia'' and ''
Genusaurus ''Genusaurus'' ( ; meaning "knee lizard") is a genus of abelisauroid dinosaur from the Early Cretaceous. Its fossils were found in France. ''Genusaurus'' is believed to have lived during the Albian stage, around 112-100 million years ago. D ...
'' which has been recovered by Tortosa ''et al.'' (2014), Baiano ''et al.'' (2020) and Salem ''et al.'' (2022). A phylogenetic analysis conducted by Zaher ''et al.'' (2020) is reproduced below. The results of an earlier analysis by Pol & Rauhut (2012) are reproduced below.


See also

*
Timeline of ceratosaur research This timeline of ceratosaur research is a chronological listing of events in the history of paleontology focused on the ceratosaurs, a group of relatively primitive, often horned, predatory theropod dinosaurs that became the apex predators of ...
* 2004 in paleontology


References


External links


Project Exploration on ''Rugops''

Chicago Park District press release on ''Rugops''
PDF Portable Document Format (PDF), standardized as ISO 32000, is a file format developed by Adobe in 1992 to present documents, including text formatting and images, in a manner independent of application software, hardware, and operating systems. ...
format
Sereno's speculation from "Science in Africa"

Artist's impressions of ''Rugops'', from Project Exploration

''Rugops''
in the Dino Directory {{Taxonbar, from=Q133025 Abelisaurids Late Cretaceous dinosaurs of Africa Cenomanian life Cretaceous Niger Fossils of Niger Fossil taxa described in 2004 Taxa named by Paul Sereno