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Moschorhinus
Moschorhinus
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Moschorhinus
Temporal range: Late Permian–Early Triassic 254–250 Ma
↓
Restoration of Moschorhinus kitchingi
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Clade: †Therocephalia
Family: †Akidnognathidae
Genus: †Moschorhinus
Broom, 1920
Species:
†M. kitchingi
Binomial name
†Moschorhinus kitchingi
Broom, 1920
Synonyms

Moschorhinus is an extinct genus of therocephalian synapsid in the family Akidnognathidae with only one species: M. kitchingi, which has been found in the Late Permian to Early Triassic of the South African Karoo Supergroup. It was a large carnivorous therapsid, reaching 1.1–1.5 metres (3.6–4.9 ft) in total body length with the largest skull comparable to that of a lion in size, and had a broad, blunt snout which bore long, straight canines.

Moschorhinus appears to have ecologically replaced the gorgonopsids as an apex predator, and hunted much like a big cat. While most abundant in the Late Permian, it survived into the Early Triassic in small numbers after the Permian Extinction, though these Triassic survivors had stunted growth.

Taxonomy

[edit]

The genus name Moschorhinus is derived from the Ancient Greek words μόσχος (mos'-khos) moschos for calf or young animal, and rhin/rhino- for nose or snout, in reference to its short, broad snout. The species name, kitchingi, refers to Mr. James Kitching, who originally found (but did not describe) the specimen.[2]

Kitching discovered the holotype specimen, a skull (best preserved, the palate), in the Karoo Supergroup in South Africa, near the village of Nieu-Bethesda. It was first described by paleontologist Robert Broom in 1920.[2] It is now one of the best known and most recognizable therapsids of the supergroup.[3]

Broom had previously named another species of therocephalian in 1907 from KwaZulu-Natal, Scymnosaurus warreni, that he later moved to Moschorhinus in 1932 as M. warreni, maintaining it as a distinct species. M. warreni was later recognised as a probable synonym of M. kitchingi by Kitching in his unpublished PhD thesis, and a re-description of the holotype in 2023 by David Groenewald and Christian Kammerer confirmed this proposal. As the older name, M. warreni would have taxonomic priority over M. kitchingi for the species. However, Groenewald and Kammerer (2023) believed it would be premature to establish M. warreni as the correct name, pending a revision of akidnognathid therocephalian taxonomy and the possibility that even older names may have seniority.[1]

The Karoo Supergroup and its outcrops

Moschorhinus remains have been found most prominently in the Upper Permian to Lower Triassic Beaufort Group.[3][4][5]

Classification

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Moschorhinus is a therocephalian, a member of the clade Eutheriodontia and the sister taxon to cynodonts and modern mammals. Moschorhinus is classified into the family Akidnognathidae, along with other large, carnivorous therapsids with strong skulls and large upper canines.[6]

Moschorhinus took over the niche once controlled by gorgonopsids. Both groups were built much like big cats. Following the extinction of Moschorhinus by the Triassic, cynodonts took over a similar niche.[6]

Description

[edit]
Side view of the head, showing the sabers and range of motion for the jaw

Moschorhinus was a large carnivore, reaching 1.1–1.5 metres (3.6–4.9 ft) in total body length and weighing 84.3 kilograms (186 lb).[7][8] The skull is similar to that of the Gorgonopsids, with large temporal fenestrae (three in total as a synapsid) and a convexly bowed palate. The skull ranged in size to comparable to a monitor lizard, to those of a lion. They possess a characteristically short, broad snout. They possess a pair of prominently long incisors, similar to the canines of saber toothed cats.[6][9]

Lateral view of Moschorhinus jaw, showing range of motion necessary for such large incisors, and upper palatal fenestrae of snout. (From van Valkenburgh and Jenkins, 2002).[6]

Snout

[edit]

The snout of Moschorhinus is characteristically short and broad. The blunt tip of the snout features a ridge running down the midline to the frontal bone.[6][4] The lower jaw is much broader than that of any other therocephalian.[6][4] The upper snout projects a bit beyond the incisors in juveniles.[4]

The nostrils were large and positioned towards the tip of the snout.[4]

Teeth

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Moschorhinus is thought to have had a dental formula of I6.C1.M3, with 6 incisors, 1 canine, and 3 postcanines in either side of the upper jaw.[2]

The incisors are housed in the premaxillae. They are large, curve slightly, and have a bell-shaped cross-section. They had smooth cutting surfaces, and, unlike those of other therocephalians, lacked facets or striae resulting from abrasion and wear.[4]

The large saber-like canines are held within the maxillae, and are quickly identifiable features of Moschorhinus. They are especially thick and strong, and uniquely circular in cross-section. In length, these sabers are comparable to gorgonopsids. While there is no real modern analogue, the most similar living example would be the clouded leopard (Neofelis nebulosa).[6]

Like other therocephalians, Moschorhinus had a reduced number of postcanines which were housed in the maxillae. In most therocephalians, the “teeth,” or rather toothlike projection (denticulations) of the pterygoid bones, are greatly reduced or missing, and in Moschorhinus they are absent.[4][10]

Skull roof

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Tracing the roof of the skull, Moschorhinus possesses small prefrontal bones above the eyes, followed by large, widened frontal bones. The parietals form a narrow sagittal crest along the midline of the skull, which houses a very basic pineal foramen.[2][4] Indentations can be seen in the temporal fossae, depressions on either side of the crest, indicating the presence of many blood vessels and nerves supplying the brain.[11]

Eye sockets

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The lacrimal bone is larger than the reduced prefrontal, and forms the majority of the eye socket. The lacrimal has a bony boss (a rounded knob) on the orbit, and a large foramen towards its inner side. The lower edge of the eye socket is formed the jugal and maxillary bones.[2] The jugal ends at the eye socket, and is not convex, as in several later therocephalians.[4]

Palate

[edit]

Overall, the palate is convex, with a broad, triangular vomer, with paired tubercles, rounded projections pointing ventrally,[6][4] similar to other akidnognathids.[2] The palatine bones (forming the back of the roof of the mouth) are enlarged and thick, especially on their outer edges where they are joined to the maxilla. On their inner edges, the palatines are joined to the pterygoid and vomer on the nose, forming part of the circumference of the nasal cavity. Between the palatine and maxilla, just behind the canines, are large foramens, presumably to allow for nerves. A slanting ridge along the middle of the palatine presumably supported a soft palate, which allowed air to travel between the nose and the lungs.[6]

The sabers require the mouth to open widely for use, making feeding difficult. The closely related Promoschorhynchus shows stiff folds (choanal crest) on the border of the nasal passage and the throat, used to keep it open and to allow for breathing while eating. The development of a secondary palate in the skull gradually evolved in therocephalians, and the choanal crest is featured in all later therocephalians.[10]

Paleobiology

[edit]
Reconstruction feeding on a Lystrosaurus

It is presumed that Moschorhinus was a cat-like predator, being able to pierce skin and hold onto struggling prey with its long canines. This is the first record of this kind of hunting technique. Given its sturdily designed, thick snout, enormous canines, and powerful jaw muscles, Moschorhinus appears to have been a daunting predator.[6]

Paleoecology

[edit]

Many vertebrate fossils have been uncovered in the Karoo Basin. Other therocephalians from the same rock level are Tetracynodon and Promoschorhynchus.[3] Moschorhinus specimens were the only large therocephalians.[12][4]

Moschorhinus seems to have gone extinct in the Early Triassic at 251 mya after the Permian Extinction by 252 mya,[13][14][15] along with 80–95% of animal species, due to a mass hypoxia event. This appears to have led to stunted growth,[3] intense seasons, reduced ecosystem diversity, and a loss of forests.[4] Fossil evidence shows that Triassic Moschorhinus grew faster than Permian ones, resulting in reduced body size in the former, largely believed to be an effect of the harsher environmental variability after the Permian Extinction (Lilliput effect).[3][16][4] Permian skulls average 207 mm (8.1 in) in length, while that of Triassic skull is only 179 mm (7.0 in).[3] Nonetheless, Triassic Moschorhinus were the largest therocephalians of their time.[3][12]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Moschorhinus

View on Grokipedia
from Grokipedia
Moschorhinus is an extinct genus of therocephalian therapsids (Synapsida: Therapsida) comprising a single species, M. kitchingi, that inhabited the Karoo Basin of South Africa during the late Permian (Changhsingian stage, Daptocephalus Assemblage Zone) and early Triassic (Induan stage, Lystrosaurus declivis Assemblage Zone). This medium-sized carnivore exhibited a stocky, robust morphology adapted for predation, with a short, strong skull up to 260 mm long, enlarged conical canines for seizing prey, and a body length of 1.1–1.3 meters, potentially weighing up to 84 kg in larger individuals. Originally described by Robert Broom in 1920 based on specimens from the Beaufort Group, Moschorhinus kitchingi belongs to the family Akidnognathidae within Eutherocephalia, a diverse of therocephalians that evolved a range of body sizes and ecological roles during the late Permian. Its postcranial skeleton, recently detailed through examination of multiple fossils, features a powerful appendicular structure including a dorsoventrally elongated , hourglass-shaped with a large deltopectoral crest, and a robust with multiple trochanters, suggesting capabilities for sustained and pursuit of prey. The vertebral column consists of approximately 34 elements (27 presacral, 3 sacral, 4 caudal), supporting a compact, muscular build suited to a macropredatory lifestyle. Ecologically, Moschorhinus occupied a to macropredator niche, coexisting with gorgonopsians in the Permian but surviving the end-Permian mass extinction—the most severe in Earth's history—unlike most contemporaries, allowing it to thrive briefly as a dominant in ecosystems. Bone histology reveals shifts in growth patterns across the extinction boundary, with pre-extinction individuals showing faster early skeletal growth rates compared to post-extinction ones, possibly reflecting environmental stressors or reduced competition. Its dental morphology, including unstriated conical canines and shearing postcanines, indicates hypercarnivory focused on prey. As one of the last large therocephalians, Moschorhinus provides critical insights into therapsid evolution, niche partitioning, and recovery dynamics following the Permo-Triassic crisis.

Discovery and etymology

History of discovery

The first fossils attributable to Moschorhinus were collected from the near in around 1910 by local collectors, with the genus receiving its formal description by Robert Broom in 1920 based on fragmentary cranial material, including an incomplete skull designated as the (NHMUK R5698). This initial material came from strata of the Beaufort Group in the Province, highlighting Moschorhinus as a prominent late Permian therocephalian predator. In 1920, Broom named the type species M. kitchingi, honoring the fossil collector James William Kitching (1896–1953), who had gathered significant specimens during the 1930s from various sites in the Karoo Basin, including the upper Permian Daptocephalus Assemblage Zone. These early finds, such as partial s from farms like Betton and Ripplemead, expanded knowledge of the taxon's distribution across the main Karoo Basin. Subsequent discoveries in the 1970s through 1990s, including multiple partial s from the Permo-Triassic boundary interval, further documented Moschorhinus as a survivor of the end-Permian extinction, with key specimens such as NMQR 3351 representing the largest known nearly complete articulated . In 2023, P. Groenewald and Christian F. Kammerer re-examined historical material, including the 1907 of Scymnosaurus warreni (now NM 188), and re-identified it as M. kitchingi based on detailed cranial comparisons that revealed overlapping diagnostic features, effectively treating M. warreni (transferred to Moschorhinus in 1932) as a junior synonym. A comprehensive postcranial study in 2024 by Michael D. Stuart and colleagues analyzed several articulated skeletons from the Karoo Basin, including NMQR 3351, revealing robust limb morphology adapted for a stocky build and confirming Moschorhinus as the largest therocephalian to persist into the .

Naming

The genus Moschorhinus was coined by the South African paleontologist Robert Broom in 1920 for a new therocephalian therapsid based on an incomplete skull from the Karoo Basin (Proceedings of the ). The name is derived from the moschos, meaning musk or calf, and rhinus, meaning nose, in reference to the broad, calf-like shape of the . The type M. kitchingi was named in honor of fossil collector James William Kitching (1896–1953), who gathered significant specimens from during .

Taxonomy

Classification

Moschorhinus is classified as a therocephalian therapsid within the family Akidnognathidae and the Eutherocephalia, part of the broader group of non-mammalian synapsids. It is closely related to other Late Permian akidnognathids such as Akidnognathus and Promoschorhynchus, with the family Akidnognathidae confirmed as monophyletic in recent phylogenetic analyses based on cranial character matrices, including synapomorphies like enlarged temporal fenestrae. Within , Akidnognathidae and related therocephalians are part of , forming a to cynodonts, with gorgonopsians positioned basal to this .

Valid species

The genus Moschorhinus is currently recognized as monotypic, containing only the valid M. kitchingi, which was originally described by Robert Broom based on a partial from the late Permian of . This is diagnosed by a combination of cranial features, including a basal length of 136–262 mm in adults, strongly posteriorly curved upper canines that are unstriated (smooth) on their labial surfaces, a short, broad rostrum, and a narrow, slit-like pineal , with minor variations in postcanine count insufficient to warrant separation at the level. A second nominal species, M. warreni (originally described as Scymnosaurus warreni in ), was proposed based on a crushed therocephalian snout from the late Permian of . However, this was re-identified as referable to Moschorhinus due to matching proportions, size, and septomaxillary morphology, and fully synonymized with M. kitchingi in 2023 owing to complete overlap in cranial measurements and absence of distinguishing diagnostic traits. Other nominal species such as M. esterhuyseni and M. natalensis are also considered likely junior synonyms of M. kitchingi. No other species have been named within the genus, and following the 2023 taxonomic revision, all late Permian and Early Triassic akidnognathid material previously assigned to Moschorhinus is attributed to M. kitchingi. This monotypic status simplifies the genus's taxonomy within Akidnognathidae and facilitates unambiguous assignment of future fossils.

Description

Size and general morphology

Moschorhinus kitchingi attained a total body length of approximately 1.1 to 1.3 meters, based on reconstructions from well-preserved specimens such as NMQR 3351. The body mass for this large individual was estimated at 84.3 kg using volumetric modeling of the skeletal elements, following established methods for synapsid mass reconstruction. The overall build of Moschorhinus was stocky and robust, featuring a broad torso and powerful limbs that supported a style of predation suited to tactics rather than prolonged pursuit. This morphology is evidenced by the particularly sturdy construction of the , , and , which provided enhanced strength for subduing prey. The tail was relatively short, contributing to the compact typical of eutherocephalian therocephalians. In terms of proportions, the skull accounted for roughly 20% of the total body , comparable to that of modern felids, with basal lengths ranging from 136 to 262 mm across known specimens. Moschorhinus adopted a quadrupedal stance, with the forelimbs exhibiting a more sprawling posture ( oriented posterolaterally) and the hind limbs approaching semi-erect ( held closer to the ).

Cranial anatomy

The of Moschorhinus exhibits a robust construction with thickened dermal bones, typical of advanced therocephalians adapted for carnivory, with adult specimens reaching basal lengths up to 260 mm. The snout is short and broad, presenting a squared-off profile that enhances structural strength for predatory functions. Each side of the upper bears six small incisors and one enlarged conical canine to facilitate prey capture. The further includes three postcanine teeth per side, characterized by serrated edges suited for shearing flesh, while the lower features complementary that enables precise occlusion and efficient biting. The roof displays a prominent , serving as a key attachment site for the jaw adductor musculature, which underscores the animal's capacity for powerful jaw closure. Large temporal fenestrae adjacent to this crest further indicate substantial development, contributing to a strong bite force. The eye sockets are prominent and forward-facing, bordered by the postorbital and jugal bones, a configuration that likely supported some degree of for accurate prey targeting. The is convex in form, featuring an incomplete secondary bony typical of advanced therocephalians. This palatal structure, combined with the overall thickening, reinforces the skull's durability against stresses encountered during hunting and feeding.

Postcranial

The postcranial of Moschorhinus kitchingi exhibits a stocky build adapted for , with robust axial and appendicular elements supporting a body length of approximately 1.1–1.3 m. The vertebral column comprises 27 presacral vertebrae, including a well-differentiated atlas-axis complex and dorsal vertebrae with robust, posterodorsally curved neural spines that provided attachment sites for epaxial musculature. Three sacral vertebrae articulate with the , while the tail is short, preserving only four caudal vertebrae in the most complete specimen, though additional caudals may have been present but not preserved. The ribcage is broad, formed by dichocephalous that are short and robust, transitioning to longer anterior and mid-dorsal ribs that shorten posteriorly, some of which are fused to their vertebrae. Uncinate processes extend from the dorsal ribs, enhancing thoracic stability and muscle support. The pectoral features a tall, narrow, and robust measuring up to 170.75 mm in height, with a convex and no distinct process; the clavicles are fan-shaped, and the interclavicle is shield-like with small lateral rami. The pelvic includes an elongated ilium with a mediolaterally thin blade up to 130 mm long, providing stability for the hindquarters, alongside robust pubis and elements featuring an . The appendicular skeleton underscores the graviportal stance of M. kitchingi, with forelimbs shorter than hindlimbs. The humerus, measuring 150–154 mm in length, bears a prominent deltopectoral crest for deltoid and pectoral muscle insertion, while the femur is more robust and longer at up to 179 mm, with three trochanters indicating strong thigh musculature. Both the manus and pes retain five digits, with short, broad metacarpals and metatarsals; the terminal phalanges are narrow and pointed, forming curved claws suited for substrate traction. Recent analysis reveals reinforced limb bones with thickened cortical bone, contributing to the overall stocky bauplan that facilitated terrestrial predation in post-Permian environments. Compared to Permian therocephalian relatives like Euchambersia, Moschorhinus displays greater robustness in the scapula, humerus, and femur, reflecting adaptations for load-bearing in a changed ecosystem.

Paleobiology

Diet and feeding mechanism

Moschorhinus was a carnivorous predator, as inferred from its , which included spatulate incisors, enlarged conical upper canines, and blade-like postcanine teeth adapted for piercing and slicing . The prominent, recurved canines, rounded in cross-section and lacking serrations or carinae, were particularly suited for stabbing and securing struggling prey during initial capture. These dental features align with those of other hypercarnivorous therocephalians, enabling efficient processing of vertebrate tissues in Permian and ecosystems. The feeding mechanism relied on a combination of cranial and postcranial adaptations for close-quarters predation. The short, robust provided mechanical leverage for powerful closure, while the postcanines facilitated shearing actions to dismember carcasses. Postcranially, Moschorhinus exhibited a stocky build with unusually robust scapulae, humeri, and femora, indicating a capacity for and rather than pursuit-based . This morphology suggests an ambush strategy, where the predator used its muscular limbs to overpower medium-sized prey, holding them in place for lethal bites. Direct evidence for diet is limited, with no preserved gut contents identified, but associated skeletal remains and the absence of specialized crushing imply a focus on soft to moderately tough prey rather than extensive consumption. Tooth wear patterns on postcanines show facets consistent with flesh-tearing and occasional contact with , supporting a hypercarnivorous niche without advanced durophagy.

Growth and

Moschorhinus displayed distinct growth patterns marked by rapid early skeletal development followed by deceleration in later ontogenetic stages, as revealed by limb histology. Juvenile and subadult bones feature woven-fibered and parallel-fibered tissues with dense, reticular vascularization indicative of fast growth rates, transitioning to less vascularized parallel-fibered and lamellar in adults greater than 80% of maximum size. Lines of arrested growth (LAGs), numbering three or more in most limb elements, suggest periodic interruptions in , likely seasonal in nature and reflecting environmental influences on development. Ontogenetic changes in Moschorhinus involved progressive robustification, with juveniles exhibiting higher relative and fewer LAGs compared to adults, which show denser cortical and more pronounced growth marks. Adults developed a stocky postcranial build supporting increased , alongside cranial modifications such as a prominent for enhanced jaw muscle attachment. Comparisons between Permian and Triassic populations highlight shifts in growth dynamics post-end-Permian extinction. Permian individuals achieved larger maximum sizes, with average skull lengths of 207 mm, while specimens averaged 179 mm, representing a Lilliput effect driven by truncated growth trajectories. Triassic bones exhibit greater vascular density, corroborating faster juvenile growth rates that may have enhanced survival under post-extinction stressors.

Paleoecology

Temporal and geographic range

Moschorhinus fossils are recorded from the Late Permian to the , encompassing the Wuchiapingian and stages (approximately 259–252 Ma) through the stage (approximately 252–251 Ma), thereby bridging the end-Permian mass extinction event. This temporal span highlights its persistence across one of the most severe biotic crises in Earth history. Geographically, all known specimens derive exclusively from the Karoo Basin in , representing deposits within the southern portion of the ancient supercontinent . Principal fossil-bearing localities are concentrated in the Beaufort Group formations, including regions near , , and . No fossils of Moschorhinus have been reported from outside this basin or elsewhere in or , underscoring its restricted paleobiogeographic distribution. In terms of , Moschorhinus is particularly abundant within the Daptocephalus Assemblage Zone of the uppermost Permian Beaufort Group and the overlying Assemblage Zone in the . It defines part of the maccaigi–Moschorhinus Subzone of the latest Permian Daptocephalus Assemblage Zone, where it co-occurs with , contributing to biostratigraphic correlations across the Permian-Triassic boundary. The fossil record shows relative scarcity in the uppermost Permian layers, suggestive of a during the interval, prior to its recovery and abundance in the .

Habitat and environment

Moschorhinus inhabited the semi-arid floodplains of the Karoo Basin in what is now during the late Permian, where seasonal rivers supported riparian forests dominated by glossopterid flora such as . These environments featured periodically wet vegetation along river channels and levees, with the climate being warm-temperate and mean annual temperatures estimated at 15–25°C based on calcite crystallization data. Fossils of Moschorhinus are primarily preserved in the mudstones, siltstones, and subordinate lenticular sandstones of the Beaufort Group, reflecting deposition in fluvial and lacustrine settings with meandering rivers and overbank fines. Following the end-Permian mass extinction, the paleoenvironment in the Karoo Basin shifted toward greater aridity, exacerbated by global warming from Siberian Traps volcanism, with evidence of fungal abundance spikes indicating widespread organic decay and ecosystem collapse. Isotopic analyses reveal a sharp rise in atmospheric CO₂ from approximately 400–800 ppmv in the latest Permian to over 2,500 ppmv in the earliest , contributing to prolonged high temperatures around 35°C and increased soil anoxia due to and habitat degradation. This transition marked a more hostile landscape with reduced vegetation cover and heightened environmental instability. The stocky postcranial morphology of Moschorhinus, including robust limbs and a compact build, likely facilitated its persistence across the Permian-Triassic boundary in these fluctuating moisture regimes, contrasting with the more gracile forms of contemporaneous therocephalians.

Role in ecosystem

In the Late Permian of the Karoo Basin, Moschorhinus occupied the of an , preying primarily on herbivorous tetrapods such as dicynodonts (e.g., Dicynodon) and pareiasaurs, as inferred from its robust macropredatory adaptations and body size comparable to modern . It competed directly with gorgonopsids for these prey resources, with subtle postcranial differences suggesting niche partitioning within the predator guild prior to the end-Permian mass extinction (EPME). evidence of bite marks on dicynodont remains from contemporaneous assemblages indicates that Moschorhinus or similar therocephalians engaged in scavenging, supplementing predation in a resource-variable environment. Following the EPME, Moschorhinus emerged as one of the few surviving large-bodied in the depauperate fauna, maintaining its role as a top predator and potentially preying on the dominant Lystrosaurus. In these recovery-phase ecosystems, it coexisted with smaller cynodonts like Thrinaxodon but filled the macropredatory niche left vacant by the of gorgonopsids, helping to regulate populations and facilitate biotic recovery through top-down control. Its relative abundance in assemblages, comprising a notable portion of fossils in sampled localities, underscores its ecological dominance during this interval. By approximately 251 Ma, Moschorhinus declined sharply, likely due to prey scarcity in the post-EPME landscape and increasing competition from emerging archosauromorphs such as Proterosuchus, which eventually replaced it as the primary large predator in communities. This turnover marked the broader shift from synapsid- to archosauromorph-dominated terrestrial ecosystems.

References

  1. https://pmc.ncbi.nlm.nih.gov/articles/PMC11326434/
  2. https://doi.org/10.7717/peerj.17765
  3. https://www.cambridge.org/core/journals/paleobiology/article/body-size-and-growth-patterns-in-the-therocephalian-moschorhinus-kitchingi-therapsida-eutheriodontia-before-and-after-the-endpermian-extinction-in-south-africa/6AE21AD5BCCB8E1DD30069F7D2F79182
  4. https://www.researchgate.net/publication/376784533_Re-identification_and_updated_stratigraphic_context_of_the_holotypes_of_the_late_Permian_tetrapods_Dicynodon_ingens_and_Scymnosaurus_warreni_from_KwaZulu-Natal
  5. https://www.gbif.org/species/4822350
  6. https://www.prehistoric-wildlife.com/species/moschorhinus/
  7. https://www.researchgate.net/publication/230029265_An_investigation_into_the_cladistic_relationships_and_monophyly_of_therocephalian_therapsids_Amniota_Synapsida
  8. https://hdl.handle.net/10539/37143
  9. https://doi.org/10.1666/12020
  10. https://archive.org/download/biostor-110301/biostor-110301.pdf
  11. https://www.biodiversitylibrary.org/part/75106
  12. https://peerj.com/articles/325/
  13. https://www.researchgate.net/publication/256305562_Body_size_and_growth_patterns_in_the_therocephalian_Moschorhinus_kitchingi_Therapsida_Eutheriodontia_before_and_after_the_end-Permian_extinction_in_South_Africa
  14. https://www.sciencedirect.com/science/article/pii/S1631068305000849
  15. https://www.researchgate.net/figure/Stratigraphy-and-geographic-distributions-of-principal-Moschorhinus-bearing-localities-in_fig1_256305562
  16. https://www.researchgate.net/publication/283841174_The_Daptocephalus_Assemblage_Zone_Lopingian_South_Africa_A_proposed_biostratigraphy_based_on_a_new_compilation_of_stratigraphic_ranges
  17. https://pubs.geoscienceworld.org/gssa/sajg/article-abstract/123/2/191/587461/Biostratigraphy-of-the-Daptocephalus-Assemblage
  18. https://pubs.geoscienceworld.org/gsa/gsabulletin/article/126/5-6/665/126011/Latest-Permian-paleosols-from-Wapadsberg-Pass
  19. https://www.giss.nasa.gov/pubs/abs/st09110u.html
  20. https://www.nature.com/articles/s41467-021-22298-7
  21. https://www.researchgate.net/publication/315869855_Warming_and_increased_aridity_during_the_earliest_Triassic_in_the_Karoo_Basin_South_Africa
  22. https://www.pnas.org/doi/10.1073/pnas.2017045118
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