Non-mammalian Synapsids Research Articles

Evolution of tooth morphological complexity and its association with the position of tooth eruption in the jaw in non-mammalian synapsids.

Heterodonty and complex molar morphology are important characteristics of mammals acquired during the evolution of early mammals from non-mammalian synapsids. Some non-mammalian synapsids had only simple, unicuspid teeth, whereas others had complex, multicuspid teeth. In this study, we reconstructed the ancestral states of tooth morphological complexity across non-mammalian synapsids to show that morphologically complex teeth evolved independently multiple times within Therapsida and that secondary simplification of tooth morphology occurred in some non-mammalian Cynodontia. In some mammals, secondary evolution of simpler teeth from complex molars has been previously reported to correlate with an anterior shift of tooth eruption position in the jaw, as evaluated by the dentition position relative to the ends of component bones used as reference points in the upper jaw. Our phylogenetic comparative analyses showed a significant correlation between an increase in tooth complexity and a posterior shift in the dentition position relative to only one of the three specific ends of component bones that we used as reference points in the upper jaw of non-mammalian synapsids. The ends of component bones depend on the shape and relative area of each bone, which appear to vary considerably among the synapsid taxa. Quantification of the dentition position along the anteroposterior axis in the overall cranium showed suggestive evidence of a correlation between an increase in tooth complexity and a posterior shift in the dentition position among non-mammalian synapsids. This correlation supports the hypothesis that a posterior shift of tooth eruption position relative to the morphogenetic fields that determine tooth form have contributed to the evolution of morphologically complex teeth in non-mammalian synapsids, if the position in the cranium represents a certain point in the morphogenetic fields.

Remarks on the functional morphology of the earliest mammals

AbstractThe earliest mammals are characterized by a series of derived characters when compared to their synapsid predecessors. In comparison to non-mammalian synapsids, these features include highly efficient teeth, a small body size, a parasagittal limb posture, as well as a reduced number of ribs and girdle elements and light-weighted tails. We argue that the mammalian body construction and its functionality are constituted by a set of partially interrelated morphological traits, including akinetic skulls, tooth anatomy, and food processing; body size, locomotor speed, and tail reduction; the posture of the body and the construction of the girdle elements; as well as body torsion and rib reduction. By discussing these features from a biomechanical view, we demonstrate that high speed was the most important evolutionary advantage of the small earliest mammals over their larger synapsid ancestors, as well as over dinosaurs at the end of the Mesozoic.

Origins of mammalian vertebral function revealed through digital bending experiments.

Unravelling the functional steps that underlie major transitions in the fossil record is a significant challenge for biologists owing to the difficulties of interpreting functional capabilities of extinct organisms. New computational modelling approaches provide exciting avenues for testing function in the fossil record. Here, we conduct digital bending experiments to reconstruct vertebral function in non-mammalian synapsids, the extinct forerunners of mammals, to provide insights into the functional underpinnings of the synapsid-mammal transition. We estimate range of motion and stiffness of intervertebral joints in eight non-mammalian synapsid species alongside a comparative sample of extant tetrapods, including salamanders, reptiles and mammals. We show that several key aspects of mammalian vertebral function evolved outside crown Mammalia. Compared to early diverging non-mammalian synapsids, cynodonts stabilized the posterior trunk against lateroflexion, while evolving axial rotation in the anterior trunk. This was later accompanied by posterior sagittal bending in crown mammals, and perhaps even therians specifically. Our data also support the prior hypothesis that functional diversification of the mammalian trunk occurred via co-option of existing morphological regions in response to changing selective demands. Thus, multiple functional and evolutionary steps underlie the origin of remarkable complexity in the mammalian backbone.

Predatory synapsid ecomorphology signals growing dynamism of late Palaeozoic terrestrial ecosystems

Terrestrial ecosystems evolved substantially through the Palaeozoic, especially the Permian, gaining much new complexity, especially among predators. Key among these predators were non-mammalian synapsids. Predator ecomorphology reflect interactions with prey and competitors, which are key controls on carnivore diversity and ecology. Therefore, carnivorous synapsids may offer insight on wider ecological evolution as the first complex, tetrapod-dominated, terrestrial ecosystems formed through the late Palaeozoic. Using morphometric and phylogenetic comparative methods, we chart carnivorous synapsid trophic morphology from the latest Carboniferous to the earliest Triassic (307-251.2 Ma). We find a major morphofunctional shift in synapsid carnivory between the early and middle Permian, via the addition of new feeding modes increasingly specialised for greater biting power or speed that captures the growing antagonism and dynamism of terrestrial tetrapod predator-prey interactions. The further evolution of new hypo- and hypercarnivorous synapsids highlight the nascent intrinsic pressures and complexification of terrestrial ecosystems across the mid-late Permian.

The fossil record of appendicular muscle evolution in Synapsida on the line to mammals: Part II-Hindlimb.

This paper is the second in a two-part series that charts the evolution of appendicular musculature along the mammalian stem lineage, drawing upon the exceptional fossil record of extinct synapsids. Here, attention is focused on muscles of the hindlimb. Although the hindlimb skeleton did not undergo as marked a transformation on the line to mammals as did the forelimb skeleton, the anatomy of extant tetrapods indicates that major changes to musculature have nonetheless occurred. To better understand these changes, this study surveyed the osteological evidence for muscular attachments in extinct mammalian and nonmammalian synapsids, two extinct amniote outgroups, and a large selection of extant mammals, saurians, and salamanders. Observations were integrated into an explicit phylogenetic framework, comprising 80 character-state complexes covering all muscles crossing the hip, knee, and ankle joints. These were coded for 33 operational taxonomic units spanning >330 Ma of tetrapod evolution, and ancestral state reconstruction was used to evaluate the sequence of muscular evolution along the stem lineage from Amniota to Theria. The evolutionary history of mammalian hindlimb musculature was complex, nonlinear, and protracted, with several instances of convergence and pulses of anatomical transformation that continued well into the crown group. Numerous traits typically regarded as characteristically "mammalian" have much greater antiquity than previously recognized, and for some traits, most synapsids are probably more reflective of the ancestral amniote condition than are extant saurians. More broadly, this study highlights the utility of the fossil record in interpreting the evolutionary appearance of distinctive anatomies.

The fossil record of appendicular muscle evolution in Synapsida on the line to mammals: Part I-Forelimb.

This paper is the first in a two-part series that charts the evolution of appendicular musculature along the mammalian stem lineage, drawing upon the exceptional fossil record of extinct synapsids. Here, attention is focused on muscles of the forelimb. Understanding forelimb muscular anatomy in extinct synapsids, and how this changed on the line to mammals, can provide important perspective for interpreting skeletal and functional evolution in this lineage, and how the diversity of forelimb functions in extant mammals arose. This study surveyed the osteological evidence for muscular attachments in extinct mammalian and nonmammalian synapsids, two extinct amniote outgroups, and a large selection of extant mammals, saurians, and salamanders. Observations were integrated into an explicit phylogenetic framework, comprising 73 character-state complexes covering all muscles crossing the shoulder, elbow, and wrist joints. These were coded for 33 operational taxonomic units spanning >330 Ma of tetrapod evolution, and ancestral state reconstruction was used to evaluate the sequence of muscular evolution along the stem lineage from Amniota to Theria. In addition to producing a comprehensive documentation of osteological evidence for muscle attachments in extinct synapsids, this work has clarified homology hypotheses across disparate taxa and helped resolve competing hypotheses of muscular anatomy in extinct species. The evolutionary history of mammalian forelimb musculature was a complex and nonlinear narrative, punctuated by multiple instances of convergence and concentrated phases of anatomical transformation. More broadly, this study highlights the great insight that a fossil-based perspective can provide for understanding the assembly of novel body plans.

Nasal turbinates of the dicynodont Kawingasaurus fossilis and the possible impact of the fossorial habitat on the evolution of endothermy.

The nasal region of the fossorial anomodont Kawingasaurus fossilis was virtually reconstructed from neutron-computed tomographic data and compared with the terrestrial species Pristerodon mackayi and other nonmammalian synapsids. The tomography of the Kawingasaurus skull reveals a pattern of maxillo-, naso-, fronto- and ethmoturbinal ridges that strongly resemble the mammalian condition. On both sides of the nasal cavity, remains of scrolled maxilloturbinals were preserved that were still partially articulated with maxilloturbinal ridges. Furthermore, possible remains of the lamina semicircularis as well as fronto- or ethmoturbinals were found. In Kawingasaurus, the maxilloturbinal ridges were longer and stronger than in Pristerodon. Except for the nasoturbinal ridges, no other ridges in the olfactory region and no remains of turbinates were recognized. This supports the hypothesis that naso-, fronto-, ethmo- and maxilloturbinals were a plesiomorphic feature of synapsids, but due to their cartilaginous nature in most taxa were, in almost all cases, not preserved. The well-developed maxilloturbinals in Kawingasaurus were probably an adaptation to hypoxia-induced hyperventilation in the fossorial habitat, maintaining the high oxygen demands of Kawingasaurus' large brain. The surface area of the respiratory turbinates in Kawingasaurus falls into the mammalian range, which suggests that they functioned as a countercurrent exchange system for thermoregulation and conditioning of the respiratory airflow. Our results suggest that the environmental conditions of the fossorial habitat led to specific sensory adaptations, accompanied by a pulse in brain evolution and of endothermy in cistecephalids, ~50 million years before the origin of endothermy in the mammalian stem line. This supports the Nocturnal Bottleneck Theory, in that we found evidence for a similar evolutionary scenario in cistecephalids as proposed for early mammals.

Revisiting the evolutionary trend toward the mammalian lower jaw in non-mammalian synapsids in a phylogenetic context.

The mammalian lower jaw comprises a single bone, the dentary, which is a unique feature among vertebrates. The lower jaws of extinct non-mammalian synapsids were composed of the dentary and several postdentary bones. Synapsid fossils exhibit variation in the dentary size relative to the overall lower jaw. An evolutionary trend toward dentary enlargement and postdentary reduction in non-mammalian synapsids has long been documented but has not been established using modern phylogenetic comparative methods. In this study, we examine the evolutionary pattern of dentary size relative to the lower jaw through phylogenetic analyses of measurements in a broad range of non-mammalian synapsid taxa. Our analyses revealed an evolutionary trend toward dentary area enlargement relative to the overall lower jaw in the lateral view across all non-mammalian synapsids. This trend is likely due to vertical expansion of the dentary given that the same trend is not evident when looking at anterior to posterior measurements of the dentary relative to the lower jaw as a whole in lateral view. Ancestral character reconstructions revealed that the evolution of the measurements was not unidirectional in non-mammalian synapsids. Our results provide no evidence of an evolutionary trend toward the dentary enlargement at the expense of postdentary bones across non-mammalian synapsids. This implies that the evolutionary origin of the mammalian lower jaw is not adequately explained by the evolutionary trend of dentary enlargement throughout non-mammalian synapsids. Instead, selection that occurred during the transition from non-mammalian cynodonts to early mammals may have produced the mammalian lower jaw.

The anteriorization of tooth position underlies the atavism of tooth morphology: Insights into the morphogenesis of mammalian molars.

The evolution and development of complex molars as a key innovation in mammals have long been of interest yet remain poorly understood. With reference to century-old theories and modern findings, we focused on the teeth of pinnipeds (Carnivora) and cetaceans (Cetartiodactyla), which are morphologically simple compared with those of other mammals, and thus can be considered a reversal toward the ancestral state of nonmammalian synapsids. By reconstructing the evolutionary history of tooth complexity for the phylogenies of Carnivora and Cetartiodactyla, we established that a secondary evolution of simple teeth from more complex molars has occurred independently multiple times. Our phylogenetic comparative analyses showed that a simplification in tooth morphology was correlated with a more anterior dentition position relative to the component bones of the upper jaw in both Carnivora and Cetartiodactyla. These results suggest that the anterior shift of tooth position relative to the morphogenetic fields present in the jaw contributed to the evolutionary simplification in molar morphology. Our findings provide insights into the developmental basis of complex mammalian dentition.

The earliest segmental sternum in a Permian synapsid and its implications for the evolution of mammalian locomotion and ventilation

The sternum is a stabilizing element in the axial skeleton of most tetrapods, closely linked with the function of the pectoral girdle of the appendicular skeleton. Modern mammals have a distinctive sternum characterized by multiple ossified segments, the origins of which are poorly understood. Although the evolution of the pectoral girdle has been extensively studied in early members of the mammalian total group (Synapsida), only limited data exist for the sternum. Ancestrally, synapsids exhibit a single sternal element and previously the earliest report of a segmental sternum in non-mammalian synapsids was in the Middle Triassic cynodont Diademodon tetragonus. Here, we describe the well-preserved sternum of a gorgonopsian, a group of sabre-toothed synapsids from the Permian. It represents an ossified, multipartite element resembling the mammalian condition. This discovery pulls back the origin of the distinctive “mammalian” sternum to the base of Theriodontia, significantly extending the temporal range of this morphology. Through a review of sternal morphology across Synapsida, we reconstruct the evolutionary history of this structure. Furthermore, we explore its role in the evolution of mammalian posture, gait, and ventilation through progressive regionalization of the postcranium as well as the posteriorization of musculature associated with mammalian breathing.

Nomenclature, comparative anatomy, and evolution of the reflected lamina of the angular in non-mammalian synapsids

ABSTRACT The reflected lamina of the angular has long been portrayed as a key component in understanding the evolution of the mammalian mandible and middle ear, yet the function of this structure in non-mammalian therapsids remains uncertain. We undertake a broad scale survey of two aspects of the reflected lamina that have been poorly characterized: the ridges and fossae on its lateral surface and the extent of the underlying angular cleft. These two features were visualized in a wide range of therapsids using CT data from the literature and blue light surface scanning. Most species exhibit a clade-specific pattern of surface topography, although dicynodonts are highly variable. A striking similarity in the general number and orientation of fossae and ridges is seen between the otherwise disparate therapsid clades, and we propose homologies in these surface features across Therapsida. These features may serve as useful phylogenetic characters in analyses of the relationships among the major therapsid clades. The angular cleft does not underlie all of what is often referred to as the ‘reflected lamina’ in the literature, and an updated definition of this term is provided. Several therapsid groups independently expanded the angular cleft to further separate the reflected lamina from the angular body. We discuss some functional consequences of our findings and note inconsistency between proposed clade-specific muscle attachment sites and the presence of homologous fossae and ridges across much of Therapsida.

Evolutionary Origins of Mammalian Axial Function Revealed through Digital Bending Experiments

The origin of mammals from non‐mammalian synapsids represents an iconic locomotor transition, often described as shift from reptile‐like sprawling limbs and extensive lateral movements of the backbone to upright limbs and sagittal backbone movements that enhance mammal specific locomotor behaviors. However, recent research in comparative anatomy has called into question the lateral‐to‐sagittal functional shift, instead suggesting that vertebral morphology in basal synapsids implies a distinct functional regime not observed in extant sprawling reptiles and represents a unique ancestral condition from which mammals evolved. To investigate this idea further, we used Autobend,a novel, experimentally validated technique for estimating vertebral osteological range of motion (oROM) from skeletons using digital modeling. We applied Autobendto seven extant mammal and reptile species and 11 exceptionally preserved non‐mammalian synapsids to estimate vertebral oROM and intervertebral joint stiffness. Results revealed a clear distinction between extant mammals and reptiles in oROM, with reptiles emphasizing lateral bending and mammals sagittal bending as expected. While most extant taxa exhibited relatively similar levels of stiffness in lateral and sagittal directions, extant lizards and salamanders displayed much more compliance in lateral bending and lower stiffness overall, in keeping with their observed axial kinematics. Conversely, most non‐mammalian synapsids displayed an intermediate condition, with stiffer intervertebral joints than extant lizards, crocodiles, and therian mammals, but similar to the pattern recovered for the modern tuatara. The mammalian condition of sagittal mobility accompanied by axial twisting in the anterior column is first observed in the crownward tritylodontid cynodont Kayentatherium. Together, these results support previous morphology‐based assertions of a distinctive ancestral vertebral function for synapsids characterized by high stiffness without the specialization for lateral bending of extant lizards or the sagittal bending and twisting observed in mammals today. This indicates a more limited role for the axial skeleton in terrestrial locomotion relative to extant groups, and a greater focus on body support than propulsion during the initial stages of synapsid evolution. This stage was followed by the evolution of mammalian‐type mobility patterns in cynodonts, and a secondary reduction in axial stiffness in certain therian clades. Given the strong association between sagittal bending and asymmetric gaits in mammals, these results provide an important first step in reconstructing the evolution of synapsid locomotor patterns.

High Blood Flow Into the Femur Indicates Elevated Aerobic Capacity in Synapsids Since the Synapsida-Sauropsida Split

Varanids are the only non-avian sauropsids that are known to approach the warm-blooded mammals in stamina. Furthermore, a much higher maximum metabolic rate (MMR) gives endotherms (including birds) higher stamina than crocodiles, turtles, and non-varanid lepidosaurs. This has led researchers to hypothesize that mammalian endothermy evolved as a second step after the acquisition of elevated MMR in non-mammalian therapsids from a plesiomorphic state of low metabolic rates. In recent amniotes, MMR correlates with the index of blood flow into the femur (Qi), which is calculated from femoral length and the cross-sectional area of the nutrient foramen. Thus,Qimay serve as an indicator of MMR range in extinct animals. Using theQiproxy and phylogenetic eigenvector maps, here we show that elevated MMRs evolved near the base of Synapsida. Non-mammalian synapsids, including caseids, edaphosaurids, sphenacodontids, dicynodonts, gorgonopsids, and non-mammalian cynodonts, showQivalues in the range of recent endotherms and varanids, suggesting that raised MMRs either evolved in synapsids shortly after the Synapsida-Sauropsida split in the Mississippian or that the low MMR of lepidosaurs and turtles is apomorphic, as has been postulated for crocodiles.

The evolution of the synapsid tusk: insights from dicynodont therapsid tusk histology.

The mammalian tusk is a unique and extreme morphotype among modern vertebrate dentitions. Tusks-defined here as ever-growing incisors or canines composed of dentine-evolved independently multiple times within mammals yet have not evolved in other extant vertebrates. This suggests that there is a feature specific to mammals that facilitates the evolution of this specialized dentition. To investigate what may underpin the evolution of tusks, we histologically sampled the tusks of dicynodont therapsids: the earliest iteration of tusk evolution and the only non-mammalian synapsid clade to have acquired such a dentition. We studied the tissue composition, attachment tissues, development and replacement in 10 dicynodont taxa and show multiple developmental pathways for the adult dentitions of dicynodont tusks and tusk-like caniniforms. In a phylogenetic context, these developmental pathways reveal an evolutionary scenario for the acquisition of an ever-growing tusk-an event that occurred convergently, but only in derived members of our sample. We propose that the evolution of an ever-growing dentition, such as a tusk, is predicated on the evolution of significantly reduced tooth replacement and a permanent soft-tissue attachment. Both of these features are fixed in the dentitions of crown-group mammals, which helps to explain why tusks are restricted to this clade among extant vertebrates.

The evolution of anteriorly directed molar occlusion in mammals

AbstractIn non-mammalian synapsids and early mammals, evolutionary transformations in the feeding and hearing apparatuses are posited to have been prerequisites for the radiation of extant mammals. Unlike most vertebrates, including many early synapsids, mammals have precise dental occlusion, a lower jaw composed of one bone, and middle ear ossicles derived from ancestral jaw bones. We illuminate a related functional transition: therian mammals (eutherians and metatherians) evolved anteriorly directed chewing strokes, which are absent in other synapsid lineages. Anteriorly directed jaw movement during occlusion necessitates anteriorly directed muscle force vectors, and we posit that a shift in muscle orientation is reflected in the fossil record by the evolutionary appearance of a posteriorly positioned angular process in cladotherians (therians and their close kin). Anteriorly directed occlusion might have been absent in earlier synapsids because of the presence of attached middle ear elements in the posterior region of the jaw that prohibited the posterior insertion of jaw musculature. These changes to the masticatory apparatus in cladotherians are likely to have permitted the evolution of novel masticatory movements, including grinding in both the anterior and medial directions (e.g. rodents and ungulates, respectively). Thus, this evolutionary transition might have been a crucial prerequisite for the dietary diversification of therians.

Phylogeny, function and ecology in the deep evolutionary history of the mammalian forelimb.

Mammals are the only living members of the larger clade Synapsida, which has a fossil record spanning 320 Ma. Despite the fact that much of the ecological diversity of mammals has been considered in the light of limb morphology, the ecological comparability of mammals to their fossil forerunners has not been critically assessed. Because of the wide use of limb morphology in testing ecomorphological hypothesis about extinct tetrapods, we sought: (i) to estimate when in synapsid history, modern mammals become analogues for predicting fossil ecologies; (ii) to document examples of ecomorphological convergence; and (iii) to compare the functional solutions of distinct synapsid radiations. We quantitatively compared the forelimb shapes of the multiple fossil synapsid radiations to a broad sample of extant Mammalia representing a variety of divergent locomotor ecologies. Our results indicate that each synapsid radiation explored different areas of morphospace and arrived at functional solutions that reflected their distinctive ancestral morphologies. This work counters the narrative of non-mammalian synapsid forelimb evolution as a linear progression towards more mammalian morphologies. Instead, a disparate array of early-evolving shapes subsequently contracted towards more mammal-like forms.

Adaptive landscapes challenge the “lateral-to-sagittal” paradigm for mammalian vertebral evolution

The evolution of mammals from their extinct forerunners, the non-mammalian synapsids, is one of the most iconic locomotor transitions in the vertebrate fossil record. In the limb skeleton, the synapsid-mammal transition is traditionally characterized by a shift from a sprawling limb posture, resembling that of extant reptiles and amphibians, to more adducted limbs, as seen in modern-day mammals. Based on proposed postural similarities between early synapsids and extant reptiles, this change is thought to be accompanied by a shift from ancestral reptile-like lateral bending to mammal-like sagittal bending ofthe vertebral column. To test this "lateral-to-sagittal" evolutionary paradigm, we used combinatorial optimization to produce functionally informed adaptive landscapes and determined the functional trade-offs associated with evolutionary changes in vertebral morphology. We show that the synapsid adaptive landscape is different from both extant reptiles and mammals, casting doubt on the reptilian model for early synapsid axial function, or indeed for the ancestral condition of amniotes more broadly. Further, the synapsid-mammal transition is characterized by not only increasing sagittal bending in theposterior column but also high stiffness and increasing axial twisting in the anterior column. Therefore,we refute the simplistic lateral-to-sagittal hypothesis and instead suggest the synapsid-mammal locomotor transition involved a more complex suite of functional changes linked to increasing regionalization of the backbone. These results highlight the importance of fossil taxa for understanding major evolutionary transitions.

Convergent dental adaptations in the serrations of hypercarnivorous synapsids and dinosaurs.

Theropod dinosaurs are well known for having a ziphodont dentition: serrated, blade-shaped teeth that they used for cutting through prey. Serrations along the carinae of theropod teeth are composed of true denticles, a complex arrangement of dentine, enamel, and interdental folds. This structure would have supported individual denticles and dissipated the stresses associated with feeding. These particular serrations were previously thought to be unique to theropod dinosaurs and some other archosaurs. Here, we identify the same denticles and interdental folds forming the cutting edges in the teeth of a Permian gorgonopsian synapsid, extending the temporal and phylogenetic distribution of this dental morphology. This remarkable instance of convergence not only represents the earliest record of this adaptation to hypercarnivory but also demonstrates that the first iteration of this feature appeared in non-mammalian synapsids. Comparisons of tooth serrations in gorgonopsians with those of earlier synapsids and hypercarnivorous mammals reveal some gorgonopsians acquired a complex tissue arrangement that differed from other synapsids.

New record of a stahleckeriid dicynodont (Therapsida, Dicynodontia) from the Late Triassic of southern Brazil and biostratigraphic remarks on the Riograndia Assemblage Zone

ABSTRACT Dicynodonts represent a speciose clade of non-mammalian synapsids that lived from the middle Permian to the Late Triassic, exhibiting a Pangaean distribution. The South American Norian fossil record is represented by two species: Jachaleria colorata from Argentina and J. candelariensis from Brazil. The Brazilian specimens come from a relatively small quarry located in the site known as Botucaraí or Botucaraí Hill (Candelária, Rio Grande do Sul). We describe here four dorsal vertebrae of a single individual referred to cf. Jachaleria candelariensis from a site named Alto Guarda Mor, located in the Faxinal do Soturno municipality, State of Rio Grande do Sul, Brazil. This taxonomic proposal and the lithological features of the site support that the outcrop belongs to the Riograndia Assemblage Zone (AZ; Candelária Sequence, Santa Maria Supersequence). Although the specimen cannot be referred unambiguously to Jachaleria candelariensis due to the lack of cranial material, it likely represents the second record of the genus in the Triassic of Brazil. We further discuss the faunal composition and relative age of some sites referred to the Riograndia AZ, which exhibit odd faunal context likely representing an AZ between the top of the Hyperodapedon AZ and the base of the Riograndia AZ.

Antiquity of “Sail-Backed” Neural Spine Hyper-Elongation in Mammal Forerunners

Neural spine hyper-elongation in tetrapods is a unique morphological adaptation that creates a dorsal sail. While this extreme morphology has appeared several times in the evolutionary history of tetrapods, it was first experimented with by the non-mammalian synapsid group known as ‘Pelycosaurs’, famously represented by the Permian apex predator Dimetrodon. Here we provide new fossil data tracing back the evolution of this morphological innovation back to the initial Carboniferous radiation of the synapsid clade. We describe a new hyper-elongated neural spine belonging to the earliest ophiacodontid synapsid Echinerpeton intermedium, from the Pennsylvanian-aged deposits of Florence, Nova Scotia. The new fossil unveils the rapid convergence and repeated evolution of the ‘dorsal sail’ morphology in Synapsida. The development of this trait may have given early synapsids a unique advantage in their early radiation over the other early amniotes, the reptiles.