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Rutiodon, one of the aquatic and superficially crocodile-like phytosaurs

Thecodontia (meaning 'socket-teeth'), now considered an obsolete taxonomic grouping, was formerly used to describe a diverse "order" of early archosaurian reptiles that first appeared in the latest Permian period and flourished until the end of the Triassic period. All of them were built somewhat like crocodiles but with shorter skulls, more erect pose and usually somewhat lighter. The group includes the ancestors of dinosaurs, pterosaurs, and crocodilians, as well as a number of extinct forms that did not give rise to any descendants. The term thecodont is still used as an anatomical description of the tooth morphology seen in these species and others.

Definition

[edit]

Thecodonts are characterized by certain shared primitive features, such as the antorbital fenestra (an opening on each side of the skull between the eye sockets and the nostrils) and teeth in sockets. The name thecodont is Greek for "socket-tooth", referring to the fact that thecodont teeth were set in sockets in the jawbones; an archosaurian characteristic that was inherited by the dinosaurs. While the taxon Thecodontia is obsolete, the term thecodont remains in use as an anatomical description of teeth in bony sockets; in addition to species formerly in this group (such as crocodiles and dinosaurs), mammals also possess thecodont dentition, which evolved independently.

They constitute an evolutionary grade of animals, a "wastebasket taxon" for any archosaur other than a crocodilian, a pterosaur, or a dinosaur (any basal archosaur). Because the cladistic paradigm only recognises monophyletic taxa as natural groups, and because thecodonts are a paraphyletic group (they include among their descendants animals that are not thecodonts), the term is no longer used as a formal name by most paleontologists, but it can still be found in older (and even fairly recent) books as a convenient shorthand for the basal archosaurs.

Taxonomic history

[edit]

Traditionally, the order Thecodontia Owen, 1859 was divided into four suborders, the Proterosuchia (early primitive forms, another paraphyletic assemblage), Phytosauria (large crocodile-like semi-aquatic animals), the Aetosauria (armoured herbivores), and the Pseudosuchia, a wastebasket taxon intended to be paraphyletic to all later archosaurs (see e.g., Alfred Sherwood Romer's Vertebrate Paleontology and Edwin H. Colbert's Evolution of the Vertebrates). Of these, only phytosaurs and aetosaurs constitute monophyletic groups, and the term Pseudosuchia was simply a catch-all term for any species that didn't fit in one of the other three sub-orders. Pseudosuchia as used in recent literature is a stem-based taxon that includes crocodile-line archosaurs, or all archosaurs (including crocodilians) that are more closely related to crocodilians than birds.

Robert Carroll, in his book Vertebrate Paleontology and Evolution (1988), replaces Pseudosuchia with Rauisuchia, Ornithosuchia (containing Ornithosuchidae and non-dinosaur, non-pterosaur Ornithodira), and the traditional category incertae sedis (of uncertain placement), while retaining the other three suborders. This is the last major textbook that still recognizes the taxon Thecodontia, as it uses a traditional Linnaean based taxonomy derived from that of Romer.

In his 1982 thesis on amniote phylogeny, Brian Gardiner attempted to define Thecodontia within a cladistic framework, containing crocodilians and haemothermata (mammals and birds), thus giving the old name to a new concept. All more recent cladistic studies (e.g., Jacques Gauthier 1986) have confirmed that Thecodontia as construed by Gardiner is a polyphyletic taxon, the members of which are not united by any shared derived characteristics. As the association of the name with the outdated concept proved to be very strong, it is now considered a historical term only, and its current usage has been abandoned.

References

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Thecodontia

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Thecodontia is an obsolete and paraphyletic taxonomic grouping in paleontology that historically encompassed a diverse assemblage of basal archosauromorph reptiles from the Late Permian through the Triassic Period (approximately 259 to 201 million years ago), distinguished by their thecodont dentition—teeth firmly anchored in deep sockets within the jawbones—and regarded as the ancestral stock giving rise to major archosaur clades, including dinosaurs, pterosaurs, birds, and crocodylomorphs.[1] These reptiles exhibited a wide range of body sizes, from small agile forms under 1 meter in length to massive predators exceeding 5 meters, and occupied varied ecological roles as apex predators, herbivores, and semi-aquatic species across Late Permian to Late Triassic deposits worldwide.[1] The group was first conceptualized in the mid-19th century by Richard Owen, who named it for the characteristic tooth implantation shared with early archosaurs, and by the early 20th century, it had become a standard "wastebasket" category for Triassic reptiles not clearly assignable to other orders, including families such as Proterosuchidae, Erythrosuchidae, Rauisuchidae, Aetosauria, and Phytosauria.[2][1] Notable genera within this historical assemblage include the basal Proterosuchus from South Africa, a crocodile-like predator; the robust sail-backed Erythrosuchus; the lightly built Euparkeria, often considered close to the dinosaur-crocodilian split; and the ornithischian-like Ornithosuchus.[2][1] With the rise of cladistic phylogenetics in the late 20th century, Thecodontia was recognized as non-monophyletic, comprising unrelated lineages united only by plesiomorphic (ancestral) traits, and was subsequently abandoned in favor of precise, hierarchy-based classifications such as Archosauriformes and Pseudosuchia.[3][1] Despite its taxonomic invalidity, the concept of thecodonts remains useful in educational contexts to illustrate the evolutionary radiation of archosaurs during the Triassic, a period marked by the recovery from the Permian-Triassic mass extinction and the eventual dominance of dinosaurs following the end-Triassic extinction event.[1]

Definition and Characteristics

Definition

The term Thecodontia was coined by the British anatomist and paleontologist Richard Owen in 1859, derived from the Greek words thḗkē (θήκη), meaning "case" or "socket," and odoús (οδούς), meaning "tooth," in reference to the distinctive mode of tooth implantation in the group's members.[4][5] Historically, Thecodontia was established as an order of reptiles primarily known from the Triassic period, characterized by thecodont dentition in which teeth are firmly anchored in deep sockets (alveoli) formed by specialized alveolar bone, allowing for secure attachment and replacement.[6] The group included primitive archosaurs that first appeared in the Late Permian and persisted until the Late Triassic, functioning as a wastebasket taxon to accommodate diverse early archosauriforms excluding the more derived lineages leading to crocodilians, dinosaurs, and pterosaurs.[7] In modern cladistic analyses, Thecodontia is regarded as paraphyletic, representing an evolutionary grade of stem-archosaurs rather than a monophyletic clade.[8]

Anatomical Features

Thecodont dentition, characterized by teeth deeply embedded in individual sockets (alveoli) within the jawbones, represents a key innovation distinguishing thecodonts from other early reptiles. This condition allowed for secure anchorage and periodic replacement through resorption of the tooth base followed by eruption of successors, enabling more efficient feeding compared to acrodont (teeth fused to the jaw margin) or pleurodont (teeth fused to the medial jaw surface) arrangements prevalent in lepidosauromorphs.[9][10] Thecodont skulls typically featured lightweight construction through the presence of an antorbital fenestra—an opening anterior to the orbit formed by the maxilla, nasal, and lacrimal bones—and a mandibular fenestra in the lower jaw, reducing overall mass while maintaining structural integrity for predatory lifestyles. Many forms exhibited elongated snouts housing the thecodont teeth, with some taxa incorporating osteoderms as dermal armor along the dorsal surface for protection against predators or environmental hazards.[11][12] In the postcranial skeleton, thecodonts displayed an erect limb posture, with limbs positioned directly beneath the body rather than sprawling laterally, facilitating more efficient locomotion and potentially higher speeds than in earlier reptiles. This was supported by a vertebral column often including two to three sacral vertebrae for robust weight distribution across the pelvis, allowing for both bipedal and quadrupedal gaits depending on the taxon.[13][14] While anatomical variations existed among thecodonts, ranging from lightly built forms adapted for agility to heavily armored ones emphasizing defense, more derived members within Archosauria exhibited archosaurian ankle configurations, including the crocodilian-type (crurotarsal) joint—where the calcaneum hinges with the fibula—or the mesotarsal joint, enabling improved terrestrial mobility across diverse habitats.[15][13]

Taxonomic History

Early Classifications

The term Thecodontia was coined by British paleontologist Richard Owen in 1859 to describe a group of Triassic reptiles characterized by their socketed (thecodont) teeth, distinguishing them from other saurians with fused or marginal dentition; this classification was based on fragmentary fossils from European localities that suggested a primitive reptilian stock.[5] Owen positioned Thecodontia as an order within Reptilia, emphasizing their biconcave vertebrae and bent trunk ribs, and included early finds from European localities that suggested a primitive reptilian stock.[5] In the late 19th century, Thecodontia was widely regarded as a basal group of archosaurs ancestral to both dinosaurs and crocodilians, serving as a morphological bridge between more primitive reptiles and the dominant Mesozoic "ruling reptiles"; key inclusions encompassed genera like Hyperodapedon, a rhynchosaur with robust dentition, and Parasuchus, an early phytosaur with elongated snouts, reflecting the diverse Triassic fauna then known from scattered global discoveries.[16] This perspective underscored Thecodontia's role in early evolutionary narratives, portraying them as versatile, crocodile-like forms that diversified rapidly after the Permian-Triassic extinction, with their socketed teeth enabling efficient predation and adaptation.[16] During the early 20th century, German paleontologist Friedrich von Huene expanded and refined Thecodontia in works from the 1910s to 1920s, dividing it into suborders such as Proterosuchia (primitive, sprawling forms like Proterosuchus), Erythrosuchia (robust carnivores like Erythrosuchus), and Rauisuchia (advanced predators with erect postures), highlighting their dominance across the Triassic as stem-archosaurs.[17] Von Huene's framework integrated new anatomical details, such as antorbital fenestrae and limb proportions, to argue for Thecodontia's centrality in archosaur evolution.[17] These classifications were profoundly shaped by major fossil discoveries, particularly from the Karoo Basin in South Africa, where early Triassic exposures yielded proterosuchians and erythrosuchians that reinforced Thecodontia's ancestral status, and the Santa Maria Formation in Brazil, where von Huene's 1920s expeditions uncovered diverse pseudosuchians, solidifying the group's perceived role as progenitors of archosaurian lineages.[17][18] By the mid-20th century, however, Thecodontia began to be viewed as obsolete under emerging cladistic methods that revealed its paraphyletic nature.

Modern Perspectives

The recognition that Thecodontia represented a paraphyletic assemblage began to emerge in the mid-20th century, notably through Alfred Romer's analysis in his 1966 edition of Vertebrate Paleontology, where he highlighted the group's heterogeneous nature and lack of shared derived traits uniting all included taxa.[19] By the 1980s, the advent of cladistic methods led to its full abandonment as a formal taxon, with Jacques Gauthier's 1986 phylogenetic analysis demonstrating that Thecodontia failed to form a monophyletic clade and instead comprised disparate lineages ancestral to various archosaur groups.[20] In contemporary taxonomy, Thecodontia is regarded as an obsolete grade or wastebasket taxon encompassing stem-group members of Archosauria, with its constituent taxa redistributed across more precise monophyletic clades such as Pseudosuchia (the crocodylomorph line), Avemetatarsalia (the bird and dinosaur line), and the former Ornithosuchia.[9] This reclassification reflects a broader shift toward phylogeny-based groupings that prioritize shared synapomorphies over evolutionary grades.[20] Ongoing debates center on the historical inclusion or exclusion of certain peripheral taxa within Thecodontia, such as proterochampsids, now placed as non-archosaurian archosauriforms, and tanystropheids, which are basal archosauromorphs outside Archosauria.[21] Additionally, the term "thecodont" persists informally in paleontology and anatomy to describe socketed (thecodont) tooth implantation in non-mammalian vertebrates, distinct from its original taxonomic connotation.[22] Twenty-first-century phylogenetic studies have further solidified these revisions through comprehensive character matrix analyses; for instance, Sterling Nesbitt's 2011 work on early archosaur evolution employed 400+ characters across 80 taxa to resolve interrelationships, emphasizing monophyletic clades like Archosauria over artificial grades such as Thecodontia.[9]

Evolutionary Significance

Origins and Diversification

Thecodontia, representing a paraphyletic assemblage of basal archosauriforms, originated in the Late Permian period, approximately 259–252 million years ago (Ma), with early representatives such as Archosaurus rossicus and Eorasaurus olsoni documented from deposits in European Russia and North America.[23] These forms emerged within the broader Archosauromorpha clade, exhibiting primitive traits like sprawling to semi-erect locomotion and versatile carnivorous feeding that likely contributed to their survival through the end-Permian mass extinction event around 252 Ma.[23] Unlike many contemporaneous tetrapods that suffered severe bottlenecks, thecodont-like archosauriforms maintained stable body sizes (e.g., pes lengths averaging 118 mm in the Permian) and persisted into the Early Triassic without significant morphological disruption, owing to their adaptability in marginal aquatic and terrestrial habitats.[23] Following the extinction, thecodonts underwent an explosive adaptive radiation during the Triassic (252–201 Ma), rapidly diversifying from small carnivorous forms into a range of ecological roles, including herbivores and semi-aquatic predators, with peak diversity occurring in the Middle to Late Triassic (Anisian–Ladinian, ~247–237 Ma).[24] This expansion filled vacant niches left by the Permian-Triassic biotic crisis, as evidenced by over 90 stem-archosaur species recorded globally, showcasing high cranial disparity for specialized diets and locomotion—such as the herbivorous rhynchosaurs and apex predatory erythrosuchids.[24] The radiation was facilitated by the supercontinent Pangea's configuration, which enabled widespread dispersal across palaeolatitudes from 0° to 55°S, and the ecological recovery of fluvial and lacustrine environments that favored their mesaxonic foot morphology and generalized dentition.[23] Environmental drivers, including the stabilization of post-extinction ecosystems and the prevalence of arid to semi-arid fluvial settings, further propelled this diversification, allowing thecodonts to achieve cosmopolitan distribution by the Early Triassic (Induan–Olenekian, ~252–247 Ma).[25] However, most thecodont lineages began declining by the Late Triassic (Carnian–Norian, ~237–208 Ma), with a sharp drop in disparity during the Carnian and near-total extinction by the Rhaetian (~208–201 Ma), as they were progressively outcompeted by more derived archosaurs, particularly early dinosaurs, in overlapping terrestrial niches.[24] This faunal turnover marked the transition from stem-archosaur dominance to the rise of crown-group archosaurs, with only crocodylomorph and avemetatarsalian lines enduring beyond the end-Triassic extinction.[25]

Relationships to Archosaurs

Thecodontia represents a paraphyletic assemblage of basal archosauromorphs that occupy stem positions relative to crown-group Archosauria, defined as the most recent common ancestor of crocodylians and avemetatarsalians (birds and their extinct relatives) and all of its descendants.[26] These taxa, spanning the Late Permian to Late Triassic, form a grade of transitional forms basal to the archosaur crown, sharing key synapomorphies such as thecodont dentition (teeth deeply socketed in the jaw bones) and a prominent fourth trochanter on the femur, which provided enhanced muscle attachment for more erect limb postures.[27] This positioning underscores Thecodontia's role as an evolutionary bridge between earlier archosauromorphs and the more derived archosaur clades. From within this stem-grade Thecodontia, two primary descendant lineages emerged early in the Triassic: Pseudosuchia, encompassing crocodylians, aetosaurs, rauisuchians, and other crocodylian-line archosaurs; and Ornithodira (synonymous with Avemetatarsalia in some analyses), including dinosaurs, pterosaurs, and birds.[26] These branches reflect an early bifurcation within Archosauria, with no evidence linking Thecodontia directly to mammalian lineages, as archosaurs are firmly nested within diapsid reptiles.[27] Phylogenetic analyses have debated the precise placement of certain "problematic" thecodonts, such as members of Euparkeriidae (e.g., Euparkeria capensis), which some studies position as stem-archosaurs close to the base of Ornithodira due to features like a gracile build and reduced palatal teeth, while others recover them outside the crown as successive outgroups to Archosauria.[28] Modern cladistic approaches, incorporating total evidence datasets with hundreds of morphological characters, have largely resolved these uncertainties by emphasizing comprehensive sampling and computational parsimony or Bayesian methods, though low support values persist for basal nodes due to incomplete fossils.[29] These debates highlight Thecodontia's paraphyletic nature as a functional grade rather than a clade, illustrating the rapid diversification and homoplasy in early archosaur evolution. The implications of Thecodontia's stem position demonstrate an early Triassic bifurcation of archosaur lines, with thecodonts serving as a diverse, transitional grade that facilitated the adaptive radiation of both pseudosuchian and ornithodiran descendants following the Permian-Triassic extinction.[26] This grade-level organization underscores the mosaic evolution of archosaur traits, such as improved terrestrial locomotion, prior to the dominance of crown groups in later Mesozoic ecosystems.

Diversity and Notable Taxa

Major Groups

Thecodontia, as historically conceived, encompassed several major subgroups of basal archosaurs that exhibited diverse adaptations during the Triassic period. These groups, including Pseudosuchia, Ornithosuchia, and more basal forms such as Proterosuchidae and Erythrosuchidae, were united by thecodont tooth implantation but later recognized as paraphyletic assemblages leading to more derived archosaurs.[27] Pseudosuchia represented the crocodile-line archosaurs within traditional Thecodontia, characterized by a crurotarsal ankle joint, extensive osteoderm armor, and pillar-like limbs supporting an erect, quadrupedal gait. This group included armored herbivores such as aetosaurs, which featured broad, beaked snouts adapted for cropping vegetation and heavy dermal plating for protection against predators. Rauisuchians, as large carnivorous forms, possessed robust skulls with ziphodont (serrated, blade-like) teeth suited for tearing flesh, along with powerful limbs for pursuing prey. Early crocodylomorphs within Pseudosuchia showed lighter builds and more agile postures, foreshadowing the diversification of modern crocodilians, with synapomorphies like ventrally constricted dorsal centra and a subhorizontal acetabulum enabling efficient terrestrial locomotion. In modern phylogenies, Ornithosuchidae—historically placed in a separate Ornithosuchia closer to the dinosaur line—are instead nested within Pseudosuchia, sharing features like the reversed crurotarsal tarsus (with socket on astragalus and peg on calcaneum).[27][30][31][32] Ornithosuchia, as originally proposed, included forms like ornithosuchids that were thought to bridge basal archosaurs and dinosaurs due to agile builds and partial bipedality. However, cladistic analyses have shown these taxa to be part of Pseudosuchia rather than transitional to ornithodirans, with traits such as reduced squamosal bones and a raised acetabulum reflecting pseudosuchian specializations rather than dinosaurian ones.[27][32][30] Other basal groups, such as Proterosuchidae and Erythrosuchidae, were historically included in Thecodontia as primitive members bridging earlier archosauromorphs and more advanced forms, though modern analyses often place them outside the strict crown-group archosaurs. Proterosuchids exhibited sprawling gaits, elongated low skulls with downturned premaxillae, and aquatic or semi-aquatic habits reminiscent of early crocodylians, with 6-9 teeth per premaxilla supporting piscivorous diets. Erythrosuchids, in contrast, were apex predators with heavily built bodies, robust skulls featuring a temporal notch, and short cervical vertebrae, achieving lengths up to 5 meters and dominating Early to Middle Triassic ecosystems through hypercarnivorous adaptations.[33][27] Intergroup variations within Thecodontia highlighted dietary and locomotor diversity, with herbivorous adaptations like the down-turned, beak-like snouts of aetosaurs contrasting sharply with the carnivorous ziphodont dentition and grasping forelimbs of rauisuchians and ornithosuchids. These differences underscored the ecological breadth of thecodonts, from armored grazers to swift hunters, while shared features like osteoderms in pseudosuchians and elevated acetabula in ornithosuchians reflected broader trends toward archosaurian specialization.[30][31]

Key Genera and Species

Proterosuchus africanus, a basal proterosuchid from the Early Triassic of South Africa, represents one of the earliest archosauromorphs with thecodont dentition. Known from nearly complete skeletons in the Lystrosaurus Assemblage Zone of the Karoo Basin, dating to around 248 million years ago, it reached lengths of 1.5 to 2 meters and exhibited a crocodile-like body plan with a long snout suited for piscivory.[33] Euparkeria capensis, a small archosauriform reptile, is known from multiple partial skeletons representing over ten individuals discovered in the Burgersdorp Formation of the Beaufort Group, near Aliwal North in the Eastern Cape of South Africa.[34] These fossils date to the early Middle Triassic, approximately 245 million years ago, during the Anisian stage.[35] Reaching about 1 meter in length, Euparkeria capensis was a gracile, likely bipedal carnivore characterized by slender limbs and a long tail, suggesting agility in terrestrial environments.[34] Its phylogenetic position as a stem archosaur close to the base of Archosauria makes it a pivotal taxon for elucidating the early radiation of archosauromorphs following the Permo-Triassic extinction.[36] Desmatosuchus smalli represents an armored pseudosuchian aetosaur, distinguished by its distinctive lateral osteoderms and cranial features from earlier congeners.[37] First identified through reexamination of specimens from the Late Triassic Chinle and Bull Canyon Formations in the southwestern United States, including Texas and Arizona, it was formally described in 2005.[37] Dating to the Norian stage around 220 million years ago, this species grew up to 4.5 meters long, exhibiting a tank-like build with prominent shoulder spikes and a herbivorous diet inferred from its dentition.[37] As a Norian-specific taxon, Desmatosuchus smalli provides biostratigraphic markers for correlating Late Triassic strata in North America.[37] Postosuchus kirkpatricki, a large rauisuchian predator, is documented from disarticulated remains of at least twelve individuals unearthed in the Dockum Group of Texas, USA.[38] These fossils, described in 1985, originate from Late Triassic deposits approximately 225 million years old.[38] Measuring 4 to 6 meters in length, it possessed a robust skull with serrated teeth and could alternate between bipedal and quadrupedal locomotion, positioning it as an apex carnivore in its ecosystem.[38] The assemblage of multiple size classes, including juveniles, hints at possible social behaviors such as pack hunting among pseudosuchians.[38] Ornithosuchus woodwardi, the namesake of Ornithosuchidae, is a Late Triassic pseudosuchian from the Lossiemouth Sandstone in Scotland, dating to the Carnian stage around 230 million years ago. Known from partial skeletons including a well-preserved skull, it measured about 3 meters in length and featured a deep skull with recurved teeth, antorbital fenestra, and a reversed crurotarsal ankle, indicating agile predatory capabilities. Modern analyses place it within Pseudosuchia, highlighting its role in early archosaur diversification.[32] Among other notable thecodontian taxa, Lagosuchus talampayensis from the Chañares Formation in northwestern Argentina exemplifies an early ornithodiran, with fossils collected during mid-20th-century expeditions led by Alfred Romer.[39] Dating to the latest Middle to earliest Late Triassic, this small, bipedal form, roughly 1 meter in length, features a slender build adapted for speed and is significant for illuminating the body plan of dinosaur precursors.[39] Erythrosuchus africanus, meanwhile, stands out as a basal archosauriform with a massively enlarged skull, known from well-preserved specimens in Early to Middle Triassic rocks of South Africa and European Russia, around 249 to 244 million years ago.[40] Reaching 4.75 to 5 meters, it served as a dominant hypercarnivorous predator in post-extinction recovery ecosystems.[40] The fossil record of thecodontians spans both Laurasia and Gondwana, with key assemblages from North American (e.g., USA), Eurasian (e.g., Russia), and southern continental (e.g., South Africa, Argentina) localities, though many specimens consist of incomplete skeletons that limit direct behavioral interpretations.

References

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  16. Basal Archosaurs: Phylogenetic Relationships and Functional ... - jstor
  17. None
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  19. Vertebrate paleontology : Romer, Alfred Sherwood, 1894
  20. Article: Saurischian monophyly and the origin of birds
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  22. The developmental origins of heterodonty and acrodonty as ...
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  26. The Early Evolution of Archosaurs: Relationships and the Origin of ...
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  29. The higher-level phylogeny of Archosauria (Tetrapoda: Diapsida)
  30. [PDF] A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding ...
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  32. Ornithosuchidae: a group of Triassic archosaurs with a unique ankle ...
  33. (PDF) 'Proterosuchia': the origin and early history of Archosauriformes
  34. New information on the braincase and inner ear of Euparkeria ...
  35. 3D hindlimb joint mobility of the stem-archosaur Euparkeria ... - Nature
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  37. A new species of the Late Triassic aetosaur Desmatosuchus ...
  38. Postosuchus, a New Thecodontian Reptile from the Triassic of Texas and the Origin of Tyrannosaurs
  39. https://ri.conicet.gov.ar/handle/11336/123975
  40. Cranial anatomy and taxonomy of the erythrosuchid archosauriform ...
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