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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 Angara Basin of Morocco.<ref name=":1">Olivier, C. 2020. Phylogeny, Paleobiogeography, and Paleophysiology of the Triassic Dicynodonts (Therapsida, Anomodontia): Contributions of the Laotian and Moroccan Forms. HAL Open Science: https://hal.science/CR2P/tel-03590277v1 </ref> Moghreberia belonged to the Stahleckeriidae family, a group of [[anomodont]] therapsids and is most commonly known by its species ''Moghreberia nmachouensis''.<ref name=":2">Kammerer, C. F., Frobisch, J., Angielczyk, K. D. 2013. On the Validity and Phylogenetic Position of Eubrachiosaurus browni, a Kannemeyeriiform Dicynodont (Anomodontia) from Triassic North America. Plos One: https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0064203</ref> 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.<ref name=":3">Moghreberia. PaleoFile: http://www.paleofile.com/Dicynodont/Moghreberia.asp</ref> 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.<ref name=":4">Corso, J.D., et al. Extinction and dawn of the modern world in the Carnian (Late Triassic). Science Advances: https://www.science.org/doi/full/10.1126/sciadv.aba0099</ref>
'''''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 Angara Basin of Morocco.{{sfn|Olivier|2020|p={{pn}}}} Moghreberia belonged to the Stahleckeriidae family, a group of [[anomodont]] therapsids and is most commonly known by its species ''Moghreberia nmachouensis''.<ref name="Kammerer Fröbisch Angielczyk 2013">{{cite journal |last1=Kammerer |first1=Christian F. |last2=Fröbisch |first2=Jörg |last3=Angielczyk |first3=Kenneth D. |title=On the Validity and Phylogenetic Position of Eubrachiosaurus browni, a Kannemeyeriiform Dicynodont (Anomodontia) from Triassic North America |journal=PLoS ONE |date=31 May 2013 |volume=8 |issue=5 |pages=e64203 |doi=10.1371/journal.pone.0064203 }}</ref> 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.<ref name=":3">{{cite web |title=Moghreberia |work=PaleoFile |url=http://www.paleofile.com/Dicynodont/Moghreberia.asp }}{{rs}}</ref> 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.<ref>{{cite journal |last1=Dal Corso |first1=Jacopo |last2=Bernardi |first2=Massimo |last3=Sun |first3=Yadong |last4=Song |first4=Haijun |last5=Seyfullah |first5=Leyla J. |last6=Preto |first6=Nereo |last7=Gianolla |first7=Piero |last8=Ruffell |first8=Alastair |last9=Kustatscher |first9=Evelyn |last10=Roghi |first10=Guido |last11=Merico |first11=Agostino |last12=Hohn |first12=Sönke |last13=Schmidt |first13=Alexander R. |last14=Marzoli |first14=Andrea |last15=Newton |first15=Robert J. |last16=Wignall |first16=Paul B. |last17=Benton |first17=Michael J. |title=Extinction and dawn of the modern world in the Carnian (Late Triassic) |journal=Science Advances |date=18 September 2020 |volume=6 |issue=38 |doi=10.1126/sciadv.aba0099 }}</ref>


==Geological and paleoenvironmental information==
==Geological and paleoenvironmental information==
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.<ref name=":1"/> The Irohalene Member is characterized by abundant vertebrate fauna remains and footprints including members of [[Dinosauromorpha]], crocodilian-stem archosaurs, phytosaurs, lepidosauromorphs and archosauromorphs <ref name=":5">Lagnaoui, A, et al. New discoveries of archosaur and other tetrapod footprints from the Timezgadiouine Formation (Irohalene Member, Upper Triassic) of the Argana Basin, western High Atlas, Morocco – Ichnotaxonomic implications. Science Direct: https://www.sciencedirect.com/science/article/abs/pii/S0031018216300359</ref> as well as actinopterygians and dipnoi.<ref name=":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.<ref name=":1"/>
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.{{sfn|Olivier|2020|p={{pn}}}} The Irohalene Member is characterized by abundant vertebrate fauna remains and footprints including members of [[Dinosauromorpha]], crocodilian-stem archosaurs, phytosaurs, lepidosauromorphs and archosauromorphs<ref>{{cite journal |last1=Lagnaoui |first1=Abdelouahed |last2=Klein |first2=Hendrik |last3=Saber |first3=Hafid |last4=Fekkak |first4=Abdelilah |last5=Belahmira |first5=Abouchouaïb |last6=Schneider |first6=Joerg W. |title=New discoveries of archosaur and other tetrapod footprints from the Timezgadiouine Formation (Irohalene Member, Upper Triassic) of the Argana Basin, western High Atlas, Morocco – Ichnotaxonomic implications |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |date=July 2016 |volume=453 |pages=1–9 |doi=10.1016/j.palaeo.2016.03.022 }}</ref> as well as actinopterygians and dipnoi.{{sfn|Olivier|2020|p={{pn}}}} 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.{{sfn|Olivier|2020|p={{pn}}}}


==Historical information and discovery==
==Historical information and discovery==
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.<ref name=":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.<ref name=":2"/> This established Moghreberia as a close relative to Stahleckeria, Ischigualastia, and Angonisaurus in addition to Placerias.<ref name=":6">Vega-Dias, C., et al. A new phylogenetic analysis of Triassic dicynodonts (Therapsida) and the
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.{{sfn|Olivier|2020|p={{pn}}}} 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.<ref name="Kammerer Fröbisch Angielczyk 2013"/> This established Moghreberia as a close relative to Stahleckeria, Ischigualastia, and Angonisaurus in addition to Placerias.<ref>{{cite journal |last1=Vega-Dias |first1=Cristina |last2=Maisch |first2=Michael W. |last3=Schultz |first3=Cesar Leandro |title=A new phyologenetic analysis of Triassic dicynodonts (Therapsida) and the systematic position of Jachaleria candelariensis from the Upper Triassic of Brazil |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |date=2 March 2004 |volume=231 |issue=2 |pages=145–166 |doi=10.1127/njgpa/231/2004/145 }}</ref> 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,<ref name="Kammerer Fröbisch Angielczyk 2013"/> whereas others claimed it was merely a synonym of Placerias and that they were one and the same.{{sfn|Olivier|2020|p={{pn}}}} 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.{{sfn|Olivier|2020|p={{pn}}}}
systematic position of Jachaleria candelariensis from the Upper Triassic
of Brazil. Research Gate: https://www.researchgate.net/profile/Michael-Maisch-2/publication/260273275_A_new_phylogenetic_analysis_of_Triassic_dicynodonts_Therapsida_and_the_systematic_position_of_Jachaleria_candelariensis_from_the_Upper_Triassic_of_Brazil/links/56f8487f08ae95e8b6d35d41/A-new-phylogenetic-analysis-of-Triassic-dicynodonts-Therapsida-and-the-systematic-position-of-Jachaleria-candelariensis-from-the-Upper-Triassic-of-Brazil.pdf</ref> 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,<ref name=":2"/> whereas others claimed it was merely a synonym of Placerias and that they were one and the same.<ref name=":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.<ref name=":1"/>


==Description and paleobiology==
==Description and paleobiology==
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===Cranial elements===
===Cranial elements===
====Skull====
====Skull====
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.<ref name=":1"/>
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.{{sfn|Olivier|2020|p={{pn}}}}

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.<ref name=":7">Cruickshank, A. R. I. 1978. Feeding Adaptations in Triassic Dicynodonts. Palaeontologia Africana: https://citeseerx.ist.psu.edu/document?repid=rep1&type=pdf&doi=bcf26b1dab432253e0646a1da5dfdce5e0651645</ref>
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.<ref name=Cruickshank1978>{{cite journal |last1=Cruickshank |first1=A. R. I. |title=Feeding adaptations in Triassic Dicynodonts |journal=Palaeontologia Africana |date=1978 |volume=21 |pages=121–132 |hdl=10539/16254 |hdl-access=free }}</ref>


====Dentition====
====Dentition====
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.<ref name=":1"/> Dicynodont tusks have been hypothesized to be for display purposes, but this is unconfirmed.<ref name=":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.<ref name=":8">Whitney, M. R., Angielczyk, K. D., Peecook, B. R., Sidor, C. A. 2021. The Evolution of the Synapsid Tusk: Insights from Dicynodont Therapsid Tusk Histology. Proceedings of the Royal Society B (Biology): https://royalsocietypublishing.org/doi/10.1098/rspb.2021.1670</ref>
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.{{sfn|Olivier|2020|p={{pn}}}} Dicynodont tusks have been hypothesized to be for display purposes, but this is unconfirmed.<ref name=Cruickshank1978/> 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.<ref>{{cite journal |last1=Whitney |first1=M. R. |last2=Angielczyk |first2=K. D. |last3=Peecook |first3=B. R. |last4=Sidor |first4=C. A. |title=The evolution of the synapsid tusk: insights from dicynodont therapsid tusk histology |journal=Proceedings of the Royal Society B: Biological Sciences |date=27 October 2021 |volume=288 |issue=1961 |doi=10.1098/rspb.2021.1670 }}</ref>


===Postcranial elements===
===Postcranial elements===
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====Vertebrae====
====Vertebrae====
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.<ref name=":1"/>
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.{{sfn|Olivier|2020|p={{pn}}}}


====Scapulacoracoid and sternum====
====Scapulacoracoid and sternum====
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.<ref name=":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.{{sfn|Olivier|2020|p={{pn}}}}


====Pelvis====
====Pelvis====
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.<ref name=":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.{{sfn|Olivier|2020|p={{pn}}}}


==Lifestyle==
==Lifestyle==
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.<ref name=":7"/>
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.<ref name=Cruickshank1978/>


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.<ref name=":9">Rey, K., Amiot, R., Fourel, F., Abdala, F., Fluteau, F., Jalil, N.E., Liu, J., Rubidge, B., Smith, R. 2017. Oxygen Isotopes Suggest Elevated Thermometabolism within Multiple Permo-Triassic Therapsid Clades. eLife Sciences: https://elifesciences.org/articles/28589</ref> 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.<ref name=":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.<ref name=":10">Olivier, C., et al. First palaeohistological inference of resting metabolic rate in an extinct synapsid, Moghreberia nmachouensis (Therapsida: Anomodontia). Biological Journal of the Linnean Society: https://academic.oup.com/biolinnean/article/121/2/409/2999304?login=false</ref>
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.<ref name="Rey Amiot Fourel et al 2017">{{cite journal |last1=Rey |first1=Kévin |last2=Amiot |first2=Romain |last3=Fourel |first3=François |last4=Abdala |first4=Fernando |last5=Fluteau |first5=Frédéric |last6=Jalil |first6=Nour-Eddine |last7=Liu |first7=Jun |last8=Rubidge |first8=Bruce S |last9=Smith |first9=Roger MH |last10=Steyer |first10=J Sébastien |last11=Viglietti |first11=Pia A |last12=Wang |first12=Xu |last13=Lécuyer |first13=Christophe |title=Oxygen isotopes suggest elevated thermometabolism within multiple Permo-Triassic therapsid clades |journal=eLife |date=18 July 2017 |volume=6 |doi=10.7554/eLife.28589 }}</ref> 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.<ref name="Rey Amiot Fourel et al 2017"/> 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.<ref>{{cite journal |last1=Olivier |first1=Chloe |last2=Houssaye |first2=Alexandra |last3=Jalil |first3=Nour-Eddine |last4=Cubo |first4=Jorge |title=First palaeohistological inference of resting metabolic rate in an extinct synapsid, Moghreberia nmachouensis (Therapsida: Anomodontia) |journal=Biological Journal of the Linnean Society |date=1 June 2017 |volume=121 |issue=2 |pages=409–419 |doi=10.1093/biolinnean/blw044 }}</ref>


== References ==
== References ==
{{Reflist}}
{{Reflist}}

==Sources==
* {{cite thesis |last1=Olivier |first1=Chloé |title=Phylogeny, paleobiogeography, and paleophysiology of the Triassic dicynodonts (Therapsida, Anomodontia) : contributions of the Laotian and Moroccan forms |date=2020 |url=https://theses.hal.science/tel-03590277 }}


== Further reading ==
== Further reading ==
* J.-M. Dutuit. 1989. ''Azarifeneria barrati'', un deuxième genre de dicynodonte du Trias supérieur marocain. Comptes rendus de l'Académie des sciences. Série 2, Mécanique, Physique, Chimie, Sciences de l'univers, Sciences de la Terre 309(2):303-306
* {{cite journal |id={{INIST|7266089}} |last1=Dutuit |first1=J.-M |title=Azarifeneria barrati, un deuxième genre de dicynodonte du Trias supérieur marocain |trans-title=Azarifeneria barrati n.g. n.sp. (Reptilia: Therapsida), second Dicynodont Genus of the Moroccan Upper Triassic |language=fr |journal=Comptes rendus de l'Académie des sciences. Série 2, Mécanique, Physique, Chimie, Sciences de l'univers, Sciences de la Terre |date=1989 |volume=309 |issue=2 |pages=303–306 }}
* Dutuit, J.-M., 1980, Principaux caracteres d'un genera de Dicynodonte du Trias marocain: Comptes rendus hebdomadaires des séances de lèAcadémie des Sciences de Paris, tomo 290, p.&nbsp;655-658.
* {{cite journal |last1=Dutuit |first1=JM |title=Principaux caracteres d'un genre de Dicynodonte du Trias marocain |trans-title=Principal characteristics of a dicynodont genus from the Moroccan Triassic |language=fr |journal=Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences, Serie D: Sciences Naturelles |date=10 March 1980 |volume=290 |issue=10 |pages=655–658 |url=https://gallica.bnf.fr/ark:/12148/bpt6k5497484p/f49.item }}
* [http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/synapsida/synapsida_index.html Classification of synapsid groups in ''Mikko's Phylogeny Archive'']
* [http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/synapsida/synapsida_index.html Classification of synapsid groups in ''Mikko's Phylogeny Archive'']


Revision as of 22:42, 23 March 2024

Moghreberia
Temporal range: Carnian
~235–222 Ma
Scientific classification Edit this 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
  • Azarifeneria Dutuit 1989

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 Angara 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]

Geological and paleoenvironmental information

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]

Historical information and discovery

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]

Description and paleobiology

Stahleckeriid mandibles
Stahleckeriid mandibles

Various examinations and analyses of both cranial and postcranial elements have outlined the characteristics displayed in Moghreberia fossils.

Cranial elements

Skull

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]

Dentition

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]

Postcranial elements

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.

Vertebrae

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]

Scapulacoracoid and sternum

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]

Pelvis

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]

Lifestyle

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]

References

  1. ^ a b c d e f g h i j k l Olivier 2020, p. [page needed].
  2. ^ a b c Kammerer, Christian F.; Fröbisch, Jörg; Angielczyk, Kenneth D. (31 May 2013). "On the Validity and Phylogenetic Position of Eubrachiosaurus browni, a Kannemeyeriiform Dicynodont (Anomodontia) from Triassic North America". PLoS ONE. 8 (5): e64203. doi:10.1371/journal.pone.0064203.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  3. ^ "Moghreberia". PaleoFile.[unreliable source?]
  4. ^ Dal Corso, Jacopo; Bernardi, Massimo; Sun, Yadong; Song, Haijun; Seyfullah, Leyla J.; Preto, Nereo; Gianolla, Piero; Ruffell, Alastair; Kustatscher, Evelyn; Roghi, Guido; Merico, Agostino; Hohn, Sönke; Schmidt, Alexander R.; Marzoli, Andrea; Newton, Robert J.; Wignall, Paul B.; Benton, Michael J. (18 September 2020). "Extinction and dawn of the modern world in the Carnian (Late Triassic)". Science Advances. 6 (38). doi:10.1126/sciadv.aba0099.
  5. ^ Lagnaoui, Abdelouahed; Klein, Hendrik; Saber, Hafid; Fekkak, Abdelilah; Belahmira, Abouchouaïb; Schneider, Joerg W. (July 2016). "New discoveries of archosaur and other tetrapod footprints from the Timezgadiouine Formation (Irohalene Member, Upper Triassic) of the Argana Basin, western High Atlas, Morocco – Ichnotaxonomic implications". Palaeogeography, Palaeoclimatology, Palaeoecology. 453: 1–9. doi:10.1016/j.palaeo.2016.03.022.
  6. ^ Vega-Dias, Cristina; Maisch, Michael W.; Schultz, Cesar Leandro (2 March 2004). "A new phyologenetic analysis of Triassic dicynodonts (Therapsida) and the systematic position of Jachaleria candelariensis from the Upper Triassic of Brazil". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 231 (2): 145–166. doi:10.1127/njgpa/231/2004/145.
  7. ^ a b c Cruickshank, A. R. I. (1978). "Feeding adaptations in Triassic Dicynodonts". Palaeontologia Africana. 21: 121–132. hdl:10539/16254.
  8. ^ Whitney, M. R.; Angielczyk, K. D.; Peecook, B. R.; Sidor, C. A. (27 October 2021). "The evolution of the synapsid tusk: insights from dicynodont therapsid tusk histology". Proceedings of the Royal Society B: Biological Sciences. 288 (1961). doi:10.1098/rspb.2021.1670.
  9. ^ a b Rey, Kévin; Amiot, Romain; Fourel, François; Abdala, Fernando; Fluteau, Frédéric; Jalil, Nour-Eddine; Liu, Jun; Rubidge, Bruce S; Smith, Roger MH; Steyer, J Sébastien; Viglietti, Pia A; Wang, Xu; Lécuyer, Christophe (18 July 2017). "Oxygen isotopes suggest elevated thermometabolism within multiple Permo-Triassic therapsid clades". eLife. 6. doi:10.7554/eLife.28589.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  10. ^ Olivier, Chloe; Houssaye, Alexandra; Jalil, Nour-Eddine; Cubo, Jorge (1 June 2017). "First palaeohistological inference of resting metabolic rate in an extinct synapsid, Moghreberia nmachouensis (Therapsida: Anomodontia)". Biological Journal of the Linnean Society. 121 (2): 409–419. doi:10.1093/biolinnean/blw044.

Sources

Further reading

  • Dutuit, J.-M (1989). "Azarifeneria barrati, un deuxième genre de dicynodonte du Trias supérieur marocain" [Azarifeneria barrati n.g. n.sp. (Reptilia: Therapsida), second Dicynodont Genus of the Moroccan Upper Triassic]. Comptes rendus de l'Académie des sciences. Série 2, Mécanique, Physique, Chimie, Sciences de l'univers, Sciences de la Terre (in French). 309 (2): 303–306. INIST 7266089.
  • Dutuit, JM (10 March 1980). "Principaux caracteres d'un genre de Dicynodonte du Trias marocain" [Principal characteristics of a dicynodont genus from the Moroccan Triassic]. Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences, Serie D: Sciences Naturelles (in French). 290 (10): 655–658.
  • Classification of synapsid groups in Mikko's Phylogeny Archive


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