Defecation in communal latrines is a common behaviour of extant mammals widely distributed among megaherbivores. This behaviour has key social functions with important biological and ecological implications. Herbivore communal latrines are only documented among mammals and their fossil record is exceptionally restricted to the late Cenozoic. Here we report the discovery of several massive coprolite associations in the Middle-Late Triassic of the Chañares Formation, Argentina, which represent fossil communal latrines based on a high areal density, small areal extension and taphonomic attributes. Several lines of evidence (size, morphology, abundance and coprofabrics) and their association with kannemeyeriiform dicynodonts indicate that these large synapsids produced the communal latrines and had a gregarious behaviour comparable to that of extant megaherbivores. This is the first evidence of megaherbivore communal latrines in non-mammal vertebrates, indicating that this mammal-type behaviour was present in distant relatives of mammals, and predates its previous oldest record by 220 Mya.
Studies of the effects of mass extinctions on ancient ecosystems have focused on changes in taxic diversity, morphological disparity, abundance, behaviour and resource availability as key determinants of group survival. Crucially, the contribution of life history traits to survival during terrestrial mass extinctions has not been investigated, despite the critical role of such traits for population viability. We use bone microstructure and body size data to investigate the palaeoecological implications of changes in life history strategies in the therapsid forerunners of mammals before and after the Permo-Triassic Mass Extinction (PTME), the most catastrophic crisis in Phanerozoic history. Our results are consistent with truncated development, shortened life expectancies, elevated mortality rates and higher extinction risks amongst post-extinction species. Various simulations of ecological dynamics indicate that an earlier onset of reproduction leading to shortened generation times could explain the persistence of therapsids in the unpredictable, resource-limited Early Triassic environments, and help explain observed body size distributions of some disaster taxa (e.g., Lystrosaurus). Our study accounts for differential survival in mammal ancestors after the PTME and provides a methodological framework for quantifying survival strategies in other vertebrates during major biotic crises.
Fossorialism is a beneficial adaptation for brooding, predator avoidance and protection from extreme climate. The abundance of fossilised burrow casts from the Early Triassic of southern Africa is viewed as a behavioural response by many tetrapods to the harsh conditions following the Permo-Triassic mass-extinction event. However, scarcity of vertebrate remains associated with these burrows leaves many ecological questions unanswered. Synchrotron scanning of a lithified burrow cast from the Early Triassic of the Karoo unveiled a unique mixed-species association: an injured temnospondyl amphibian (Broomistega) that sheltered in a burrow occupied by an aestivating therapsid (Thrinaxodon). The discovery of this rare rhinesuchid represents the first occurrence in the fossil record of a temnospondyl in a burrow. The amphibian skeleton shows signs of a crushing trauma with partially healed fractures on several consecutive ribs. The presence of a relatively large intruder in what is interpreted to be a Thrinaxodon burrow implies that the therapsid tolerated the amphibian’s presence. Among possible explanations for such unlikely cohabitation, Thrinaxodon aestivation is most plausible, an interpretation supported by the numerous Thrinaxodon specimens fossilised in curled-up postures. Recent advances in synchrotron imaging have enabled visualization of the contents of burrow casts, thus providing a novel tool to elucidate not only anatomy but also ecology and biology of ancient tetrapods.
Secondary adaptation to aquatic life occurred independently in several amniote lineages, including reptiles during the Mesozoic and mammals during the Cenozoic. These evolutionary shifts to aquatic environments imply major morphological modifications, especially of the feeding apparatus. Mesozoic (250-65 Myr) marine reptiles, such as ichthyosaurs, plesiosaurs, mosasaurid squamates, crocodiles, and turtles, exhibit a wide range of adaptations to aquatic feeding and a broad overlap of their tooth morphospaces with those of Cenozoic marine mammals. However, despite these multiple feeding behavior convergences, suction feeding, though being a common feeding strategy in aquatic vertebrates and in marine mammals in particular, has been extremely rarely reported for Mesozoic marine reptiles.
Based on specimens previously identified as Tropidostoma, a new taxon of dicynodont (Bulbasaurus phylloxyron gen. et sp. nov.) from the Karoo Basin of South Africa is described. Bulbasaurus is a medium-sized dicynodont (maximum dorsal skull length 16.0 cm) restricted to the Tropidostoma Assemblage Zone (early Lopingian) of the Beaufort Group. Bulbasaurus can be distinguished from Tropidostoma by an array of characters including the presence of a tall, sharp premaxillary ridge, large, rugose, nearly-confluent nasal bosses, a nasofrontal ridge, massive tusks, robust pterygoids, prominently twisted subtemporal bar, and absence of a distinct postfrontal. Inclusion of Bulbasaurus in a phylogenetic analysis of anomodont therapsids recovers it as a member of Geikiidae, a clade of otherwise later Permian dicynodonts such as Aulacephalodon and Pelanomodon. Bulbasaurus exhibits many of the characters typical of adult Aulacephalodon, but at substantially smaller skull size (these characters are absent in comparably-sized Aulacephalodon juveniles), suggesting that the evolution of typical geikiid morphology preceded gigantism in the clade. Bulbasaurus is the earliest known geikiid and the only member of the group known from the Tropidostoma Assemblage Zone; discovery of this taxon shortens a perplexing ghost lineage and indicates that abundant clades from the later Permian of South Africa (e.g., Geikiidae, Dicynodontoidea) may have originated as rare components of earlier Karoo assemblage zones.
Many lizards can drop a portion of their tail in response to an attack by a predator, a behaviour known as caudal autotomy. The capacity for intravertebral autotomy among modern reptiles suggests that it evolved in the lepidosaur branch of reptilian evolution, because no such vertebral features are known in turtles or crocodilians. Here we present the first detailed evidence of the oldest known case of caudal autotomy, found only among members of the Early Permian captorhinids, a group of ancient reptiles that diversified extensively and gained a near global distribution before the end-Permian mass extinction event of the Palaeozoic. Histological and SEM evidence show that these early reptiles were the first amniotes that could autotomize their tails, likely as an anti-predatory behaviour. As in modern iguanid lizards, smaller captorhinids were able to drop their tails as juveniles, presumably as a mechanism to evade a predator, whereas larger individuals may have gradually lost this ability. Caudal autotomy in captorhinid reptiles highlights the antiquity of this anti-predator behaviour in a small member of a terrestrial community composed predominantly of larger amphibian and synapsid predators.
Birds, dinosaurs, crocodilians, pterosaurs and their close relatives form the highly diverse clade Archosauriformes. Archosauriforms have a deep evolutionary history, originating in the late Permian, prior to the end-Permian mass extinction, and radiating in the Triassic to dominate Mesozoic ecosystems. However, the origins of this clade and its extraordinarily successful body plan remain obscure. Here, we describe an exceptionally preserved fossil skull from the Lower Triassic of Brazil, representing a new species, Teyujagua paradoxa, transitional in morphology between archosauriforms and more primitive reptiles. This skull reveals for the first time the mosaic assembly of key features of the archosauriform skull, including the antorbital and mandibular fenestrae, serrated teeth, and closed lower temporal bar. Phylogenetic analysis recovers Teyujagua as the sister taxon to Archosauriformes, and is congruent with a two-phase model of early archosauriform evolution, in response to two mass extinctions occurring at the end of the Guadalupian and the Permian.
The incompleteness of the fossil record obscures the origin of many of the more derived clades of vertebrates. One such group is the Ichthyopterygia, a clade of obligatory marine reptiles that appeared in the Early Triassic epoch, without any known intermediates. Here we describe a basal ichthyosauriform from the upper Lower Triassic (about 248 million years ago) of China, whose primitive skeleton indicates possible amphibious habits. It is smaller than ichthyopterygians and had unusually large flippers that probably allowed limited terrestrial locomotion. It also retained characteristics of terrestrial diapsid reptiles, including a short snout and body trunk. Unlike more-derived ichthyosauriforms, it was probably a suction feeder. The new species supports the sister-group relationships between ichthyosauriforms and Hupehsuchia, the two forming the Ichthyosauromorpha. Basal ichthyosauromorphs are known exclusively from south China, suggesting that the clade originated in the region, which formed a warm and humid tropical archipelago in the Early Triassic. The oldest unequivocal record of a sauropterygian is also from the same stratigraphic unit of the region.
Nocturnality is a key evolutionary innovation of mammals that enables mammals to occupy relatively empty nocturnal niches. Invasion of ancestral mammals into nocturnality has long been inferred from the phylogenetic relationships of crown Mammalia, which is primarily nocturnal, and crown Reptilia, which is primarily diurnal, although molecular evidence for this is lacking. Here we used phylogenetic analyses of the vision genes involved in the phototransduction pathway to predict the diel activity patterns of ancestral mammals and reptiles. Our results demonstrated that the common ancestor of the extant Mammalia was dominated by positive selection for dim-light vision, supporting the predominate nocturnality of the ancestral mammals. Further analyses showed that the nocturnality of the ancestral mammals was probably derived from the predominate diurnality of the ancestral amniotes, which featured strong positive selection for bright-light vision. Like the ancestral amniotes, the common ancestor of the extant reptiles and various taxa in Squamata, one of the main competitors of the temporal niches of the ancestral mammals, were found to be predominate diurnality as well. Despite this relatively apparent temporal niche partitioning between ancestral mammals and the relevant reptiles, our results suggested partial overlap of their temporal niches during crepuscular periods.
The early tetrapod Acanthostega gunnari is an iconic fossil taxon exhibiting skeletal morphology reflecting the transition of vertebrates from water onto land. Computed tomography data of two Acanthostega skulls was segmented using visualization software to digitally separate bone from matrix and individual bones of the skull from each other. A revised description of cranial and lower jaw anatomy in this taxon based on CT data includes new details of sutural morphology, the previously undescribed quadrate and articular bones, and the mandibular symphysis. Sutural morphology is used to infer loading regime in the skull during feeding, and suggests Acanthostega used its anterior jaws to initially seize prey while smaller posterior teeth were used to restrain struggling prey during ingestion. Novel methods were used to repair and retrodeform the skull, resulting in a three-dimensional digital reconstruction that features a longer postorbital region and more strongly hooked anterior lower jaw than previous attempts while supporting the presence of a midline gap between the nasals and median rostrals.