Moghreberia Temporal range: Carnian ~ | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Dicynodontia |
Family: | † Stahleckeriidae |
Subfamily: | † Placeriinae |
Genus: | † Moghreberia Dutuit 1980 |
Type species | |
Moghreberia nmachouensis Dutuit 1980 | |
Synonyms | |
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Moghreberia is an extinct genus of dicynodont predicted to have lived only in the mid-Triassic, primarily during the early middle Carnian and found only in the Argana Basin of Morocco. [1] Moghreberia belonged to the Stahleckeriidae family, a group of anomodont therapsids and is most commonly known by its species Moghreberia nmachouensis. [2] Its name is derived from the Arabic phrase al-Maghrib al-Aqsa meaning “the far west”, a term used by Arabic scholars to refer to the approximate region of Morocco, the area in which this animal’s fossil was first discovered. [3] The extinction of many dicynodonts has been attributed to pressures of the Carnian Pluvial Episode, which occurred around 234-232 Ma and generated major ecological and climate changes for years to come. [4]
The first remains of Moghreberia included two poorly preserved skulls discovered in the Irohalene Member, a section of the Timezgadiouine Formation in the Argana Basin of modern-day Morocco. [1] The Irohalene Member is characterized by abundant vertebrate fauna remains and footprints including members of Dinosauromorpha, crocodilian-stem archosaurs, phytosaurs, lepidosauromorphs and archosauromorphs [5] as well as actinopterygians and dipnoi. [1] The Argana Basin is in the western part of the High Atlas of Morocco and consists of three distinct formations known as the Timezgadiouine, Ikakern and the Bigoudine formations. Like Moghreberia nmachouensis, most dicynodont remains in this region were found in the Timezgadiouine formation. Geology of this formation suggests a dry and humid climate with shallow, still-bodied water. [1]
After discovery of the initial two skulls in the Argana Basin, these remains were first detailed, and the genus and species named, by JM Dutuit in 1980, in which he emphasized similarities to other kannemeyerids. Dutuit further detailed the genus using more well-preserved partial cranial remains discovered in the Argana Basin as well in 1988. It was using this new fossil that he initially categorized Moghreberia into Kannemeyeridae. Dutuit did, however, recognize similarities between Moghreberia and the North American genus Placerias within the family Stahleckeriidae. [1] This observation was later supported by computer-assisted phylogenetic analysis by Kammerer et al. which concluded Moghreberia was most closely related to Placerias, placing it in Stahleckeriidae as opposed to Kannemeyeridae. [2] This established Moghreberia as a close relative to Stahleckeria, Ischigualastia, and Angonisaurus in addition to Placerias. [6] Based on these few isolated cranial elements and the complete lack of postcranial remains, the taxonomic validity of Moghreberia was highly debated. Some authors claimed it was a sister taxon of Placerias, as previously mentioned, [2] whereas others claimed it was merely a synonym of Placerias and that they were one and the same. [1] It wasn’t until 2020 that its taxonomic validity was reanalyzed and the gap in phylogenetic positioning closed with the first postcranial analysis of Moghreberia nmachouensis, which was made possible due to the discovery of a nearly complete skeleton of the individual. This analysis concluded that Moghreberia was in fact phylogenetically unique and is likely more closely related to Lisowicia of Poland than to North American Placerias. [1]
Various examinations and analyses of both cranial and postcranial elements have outlined the characteristics displayed in Moghreberia fossils.
In the initial primary analysis of Moghreberia skulls, Dutuit listed multiple unique features of Moghreberia cranial elements helping to distinguish it from other dicynodonts. This genus was characterized by a low, dorsoventrally, and laterally expanded occipital surface as well as primarily horizontal lateral mandibular branches. Dutuit also noted a highly pointed tip of the snout and a dorsal margin of the erupted portion of the canine tusk anterior to the nasal cavity. The skull overall was relatively large and narrow for a dicynodont, measuring over 40 cm. Small lateral expansion of the squamosals is evident. This genus can also be characterized by a highly angled intertemporal bar, elongate posterior parietal processes, and a deeply depressed preparietal. Two cranial elements helping to distinguish Moghreberia from Placerias include a lack of caniniform depression and elongate postnarial excavation. [1]
Being of the Stahleckeriidae family, Moghreberia exhibits a blunt snout as opposed to the rounded snouts of Kennemeyeriidae. The former shape, the one seen in Moghreberia, was more common among dicynodonts than the latter. [7]
Despite the fact that the family, Stahleckeriidae, is defined as tuskless, Moghreberia, like its close relative Placerias, did in fact have large tusks protruding anterior to the naval cavity on the skull. In fact, Moghreberia had a complete absence of teeth besides these paired maxillary tusks. [1] Dicynodont tusks have been hypothesized to be for display purposes, but this is unconfirmed. [7] Being a Triassic dicynodont implies that these were likely ever-growing and composed of thick dentine walls, an enamel cap and roots growing deep into the maxilla. Larger individuals, like Moghreberia, exhibit narrowing of the pulp cavity as well. [8]
With the discovery of a nearly complete skeleton of Moghreberia, postcranial elements could finally be observed and were officially detailed in 2020 by Chloe Olivier. Moghreberia was overall a large dicynodont.
The vertebrae of Moghreberia were determined to be amphicoelous being both antiorly and posteriorly concave. Due to decreased obtuse angles between the zygopophyses of the dorsal vertebrae, this genus would be less flexible in lateral movement. It has also been confirmed that Moghreberia has five sacral vertebrae accompanied by five sacral ribs. Due to the large size difference between the first and fifth caudal vertebrae, it is predicted that caudal vertebrae size rapidly decreases towards the posterior. [1]
The scapula of the specimen is elongate with enlarged dorsal and even more enlarged ventral regions. The precoracoid and coracoid are anteriorly fused with the scapula. A coracoid foramen is clearly noticeable. The interclavicle is characterized by a triangular anterior region and a longer posterior region. The sternum exhibits sternal bosses presenting an anterodorsal rounded surface. Moghreberia, being a member of Placeriinae, displayed a distinct posterior projection on the posterior surface of the scapula. [1]
An isolated, unfused ilium, ischium and pubis indicate a very flexible pelvis in Moghreberia. The articular process of the ischia is described as stout and ventrally constricted. The pubic tubercle is made of a thick process extending from the pubis anteroventrally. [1]
Members of Stahlekeriidae, like all dicynodonts, were herbivorous. It is postulated that Stahlekeriidae were also browsers, as opposed to grazers, feeding at head height. The characteristic broad snout of Moghreberia and other Stahlekeriidae implies a lack of precision and selectivity in feeding that was possible with the narrow snouts of Kannemeyeridae. The tusks of Moghreberia are thought to serve display purposes rather than any feeding purposes. [7]
It has also been hypothesized that Moghreberia was endothermic, making it one of the earliest endotherms discovered. Studies by Rey et al. focused on oxygen isotopes and fibrolamellar bone to make this determination. Oxygen occurs in two isotopes in the tissue and is temperature dependent. Oxygen isotope 18 occurs in greater amounts in greater body temperatures. Moghreberia was found to have high relative levels of this isotope, implying increased body temperature and increased metabolic activity. [9] Additionally, fibrolamellar bone can be used as evidence of sustained fast growth. Moghreberia was found to possess fibrolamellar bone, further suggesting a relatively high metabolic rate able to support endothermy. [9] Olivier conducted a similar analysis of fibrolamellar bone in the femur and humerus and again found relatively high levels which were used to further compute relatively high metabolic rates. [10]
Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typically toothless beak, unique amongst all synapsids. Dicynodonts first appeared in Southern Pangaea during the mid-Permian, ca. 270–260 million years ago, and became globally distributed and the dominant herbivorous animals in the Late Permian, ca. 260–252 Mya. They were devastated by the end-Permian Extinction that wiped out most other therapsids ca. 252 Mya. They rebounded during the Triassic but died out towards the end of that period. They were the most successful and diverse of the non-mammalian therapsids, with over 70 genera known, varying from rat-sized burrowers to elephant-sized browsers.
Placerias is an extinct genus of dicynodonts that lived during the Carnian to the Norian age of the Triassic Period. Placerias belongs to a group of dicynodonts called Kannemeyeriiformes, which was the last known group of dicynodonts before the taxon became extinct at the end of the Triassic.
Diictodon is an extinct genus of pylaecephalid dicynodont that lived during the Late Permian period, approximately 255 million years ago. Fossils have been found in the Cistecephalus Assemblage Zone of the Madumabisa Mudstone of the Luangwa Basin in Zambia and the Tropidostoma Assemblage Zone of the Teekloof Formation, Tapinocephalus Assemblage Zone of the Abrahamskraal Formation, Dicynodon Assemblage Zone of the Balfour Formation, Cistecephalus Assemblage Zone of the Middleton or Balfour Formation of South Africa and the Guodikeng Formation of China. Roughly half of all Permian vertebrate specimens found in South Africa are those of Diictodon. This small herbivorous animal was one of the most successful synapsids in the Permian period.
Azendohsaurus is an extinct genus of herbivorous archosauromorph reptile from roughly the late Middle to early Late Triassic Period of Morocco and Madagascar. The type species, Azendohsaurus laaroussii, was described and named by Jean-Michel Dutuit in 1972 based on partial jaw fragments and some teeth from Morocco. A second species from Madagascar, A. madagaskarensis, was first described in 2010 by John J. Flynn and colleagues from a multitude of specimens representing almost the entire skeleton. The generic name "Azendoh lizard" is for the village of Azendoh, a local village near where it was first discovered in the Atlas Mountains. It was a bulky quadruped that unlike other early archosauromorphs had a relatively short tail and robust limbs that were held in an odd mix of sprawled hind limbs and raised forelimbs. It had a long neck and a proportionately small head with remarkably sauropod-like jaws and teeth.
Ischigualastia is an extinct genus of large dicynodont therapsids that lived during the Late Carnian age and the Early Norian age of the Late Triassic Period. The genus was found in and named after the Ischigualasto Formation of the Ischigualasto-Villa Unión Basin in northwestern Argentina. It has been placed in the family Stahleckeriidae.
Eodicynodon is an extinct genus of dicynodont therapsids, a highly diverse group of herbivorous synapsids that were widespread during the middle-late Permian and early Triassic. As its name suggests, Eodicynodon is the oldest and most primitive dicynodont yet identified, ranging from the middle to late Permian and possessing a mix of ancestral anomodont/therapsid features and derived dicynodont synapomorphies.
Wadiasaurus is an extinct genus of dicynodont from the family Kannemeyeria, that lived in herds from the early to Middle Triassic. Substantial fossorial evidence of W. indicus was recovered from Yerrapalli Formation of the Pranhita-Godavari valley, India, and it is so far the only Kannemeyeriid known for certain from India. The Kannemeyeriiformes underwent a significant diversification during the middle Triassic, with roughly 40 known species distributed worldwide. All Kannemeyeriiformes were medium to large bodied, graviportal herbivores with relatively erect posture and gait. Wadiasaurus indicus is currently the only known species of Wadiasaurus.
Myosaurus is a genus of dicynodont synapsids. Myosaurus was a small, herbivorous synapsid that existed around the early Triassic period. All of the fossils found of this species were found in Antarctica and South Africa. Compared to other fossils found from species that existed during this time, the Myosaurus is not common in the fossil record. This is due to a shortage of discovered fossils that possess characteristics unique to the Myosaurus. Notably, under 130 fossil fragments have been found that have been classified as Myosauridae, and almost all have been skulls. These skulls can be classified as Myosaurus because this species, unlike other dicynodonts, do not possess tusks or postfrontal teeth. The only species identified in the family Myosauridae is the Myosaurus gracilis, or M. gracilis. It should be recognized that the Myosaurus is almost always referred to as the M. gracilis in scientific research.
Biseridens is an extinct genus of anomodont therapsid, and one of the most basal anomodont genera known. Originally known from a partial skull misidentified as an eotitanosuchian in 1997, another well-preserved skull was found in the Qingtoushan Formation in the Qilian Mountains of Gansu, China, in 2009 that clarified its relationships to anomodonts, such as the dicynodonts.
Dicynodontoides is a genus of small to medium-bodied, herbivorous, emydopoid dicynodonts from the Late Permian. The name Dicynodontoides references its “dicynodont-like” appearance due to the caniniform tusks featured by most members of this infraorder. Kingoria, a junior synonym, has been used more widely in the literature than the more obscure Dicynodontoides, which is similar-sounding to another distantly related genus of dicynodont, Dicynodon. Two species are recognized: D. recurvidens from South Africa, and D. nowacki from Tanzania.
Eosimops is an extinct genus of pylaecephalid dicynodonts. They were small synapsids superficially resembling modern mammals. Eosimops is known from several skull specimens, as well as one complete skeleton. Eosimops lived during the Middle Permian of South Africa.
Elephantosaurus is an extinct genus of dicynodont from the Middle Triassic (Ladinian) Bukobay Formation. The holotype and only known specimen, catalogued as PIN 525/25, is a fragment of the skull that includes portions of the left interorbital region and nasal bones, and suggests a very large animal with a skull at least 30 centimetres (12 in) wide. The bones of the skull roof are also unusually thick. While usually considered a member of the Stahleckeriidae, generally due to its size, it probably falls just outside the group due to its frontal bone contributing substantially to the margin of the eye socket.
Kawingasaurus is an extinct genus of dicynodont therapsid from the Late Permian Usili Formation of Tanzania. It is a member of the family Cistecephalidae, and like other cistecephalids it is thought to have been fossorial. It is a member of the family Cistecephalidae. Cistephalidae includes genera Cisteceohalus, Cistecephaloides and Kawingasaurus. Greek for Saurus meaning “lizard” appears as a suffix denoting a reptilian origin. Living between 254.17 and 259.9 million years ago in the late Permian and believed to have the first and last recorded appearance in this time period. It lived in deep burrows as a suggested by most burrowing dicynodonts from evaluation of cranial sutures, vestibule inflation and enlarged stapes foot plates which are thought to be functionally correlated with bone-conduction hearing; all observed in fossorial vertebrates which use seismic signals as communication.
Odontocyclops is an extinct genus of Dicynodonts that lived in the Late Permian. Dicynodonts are believed to be the first major assemblage of terrestrial herbivores. Fossils of Odontocyclops have been found in the Karoo Basin of South Africa and the Luangwa Valley of Zambia. The phylogenetic classification of Odontocyclops has been long under debate, but most current research places them as their own genus of Dicynodonts and being very closely related to Rhachiocephalus and Oudenodon.
Pelanomodon is an extinct genus of dicynodont therapsids that lived in the Late Permian period. Fossil evidence of this genus is principally found in the Karoo Basin of South Africa, in the Dicynodon Assemblage Zone. Lack of fossil record after the Late Permian epoch suggests that Pelanomodon fell victim to the Permian-Triassic extinction event.
Tetragonias is an extinct genus of dicynodont from the Anisian Manda Beds of Tanzania. With tetra meaning “four,” and goni meaning “angle,” the name references the square shape of the Tetragonias skull when viewed dorsally. Not to be confused with the plant Tetragonia,Tetragonias were dicynodont anomodonts discovered in the late 1960s by paleontologist A. R. I. Cruickshank in the Manda Formation. Only the type species, T. njalilus, has been recognized.
Lisowicia is an extinct genus of giant dicynodont synapsid that lived in what is now Poland during the late Norian or earliest Rhaetian age of the Late Triassic Period, about 210–205 million years ago. Lisowicia is the largest known dicynodont, as well as the largest non-mammalian synapsid, reaching about 4.5 metres (15 ft) long, standing up to 2.6 metres (8.5 ft) tall at the hips and weighing around 5–7 metric tons, comparable in size to modern elephants. It was also one of the last dicynodonts, living shortly before their extinction at the end of the Triassic period. Fossils of a giant dicynodont were known from Poland since 2008, but Lisowicia was not named and officially described as a new species until late 2018.
Taoheodon is an extinct genus of dicynodont therapsid from the Sunjiagou Formation in the Shanxi province of China, dated to the Wuchiapingian age of the Late Permian. Its type and only known species is T. baizhijuni. Taoheodon was a close relative of the well known Dicynodon, and may represent a biogeographical link between the South African Dicynodon and similar dicynodonts found in Laos.
The Timezgadiouine Formation, sometimes spelled as the Timesgadiouine Formation, is a Triassic geological formation in the Argana Basin of Morocco. It is a succession of red bed sediments spanning from the Olenekian to at least the Carnian, encompassing members T3 to T5 of the Argana Group. It is preceded by the Permian Ikakern Formation and succeeded by the Late Triassic Bigoudine Formation.
Argodicynodon is an extinct genus of stahleckeriid dicynodont from the Late Triassic (Norian) of Texas in the United States. The type and only known species A. boreni was named in 2023 by palaeontologists Bill Mueller and colleagues from fossils collected from 1993 to 2014. The combined name is translated as "Boren's swift dicynodont" from the Ancient Greek argos It was discovered in the Boren Quarry in the Tecovas Formation of Texas, strata which has also been referred to as the lower Cooper Canyon Formation, and is known from isolated remains of multiple individuals representing the skull, mandibles, vertebrae, pectoral girdle, forelimb and pelvic girdle. The holotype specimen is a partial skull missing the front of the snout, palate and jaw joints, with a total length estimated to be 33 centimetres (13 in) long. A mandible from a larger individual is estimated to correspond to a skull 49 centimetres (19 in) long. Argodicynodon was related to and resembled the well-known Placerias, but had a tall, narrow sagittal crest rising sharply from behind the eyes instead of a broad flat intertemporal region. Unlike Placerias, Argodicynodon has prominent and exposed, but slender, tusks, more similar to the related Moroccan placeriine Moghreberia. Argodicynodon is also distinguished from Placerias by the arrangement of the joints between the bones of the skull, particularly of the roof of the skull along the sagittal crest. Phylogenetic analyses corroborated the placeriine identity of Argodicynodon.