No abstract

As paleontological studies are generally distorted by gaps and biases in the fossil record, it is important to assess its completeness. Here we address the fossil record of Parareptilia, a Permian–Triassic amniote clade, applying two measures of specimen completeness: the skeletal completeness metric (SCM) and the character completeness metric (CCM). The SCM quantifies how much of the skeletal material of a taxon is preserved, whereas the CCM measures the amount of phylogenetic information available. The latter was implemented using two different approaches. In this study, we compare three completeness metrics. Two CCM implementations show a strong correlation with each other, but only the second implementation of the CCM correlates significantly with the SCM, possibly due to character selection in phylogenetic data sets. There is no correlation between diversity of parareptiles and their completeness, implying that the observed fluctuations in diversity are not driven by the completeness of the fossils. The mean completeness of parareptiles through time is consistently high compared with previously studied tetrapod clades, suggesting that most parareptile taxa are based on reasonably complete specimens. Clade-specific differences reveal no link between body size and completeness. However, the analyses confirm the impact of ecology, with aquatic mesosaurids being better preserved than terrestrial taxa.

The development of the vertebral column has been studied extensively in modern amniotes, yet many aspects of its evolutionary history remain enigmatic. Here we expand the existing data on four major vertebral developmental patterns in amniotes based on exceptionally well-preserved specimens of the early Permian mesosaurid reptile Stereosternum: (i) centrum ossification, (ii) neural arch ossification, (iii) neural arch fusion, and (iv) neurocentral fusion. We retrace the evolutionary history of each pattern and reconstruct the ancestral condition in amniotes. Despite 300 million years of evolutionary history, vertebral development patterns show a surprisingly stability in amniotes since their common ancestor. We propose that this conservatism may be linked to constraints posed by underlying developmental processes across amniotes. However, we also point out that mammals and birds differ more strongly from the ancestral condition than other clades, which might be linked to a stronger regionalization of the column in these two clades.

Abstract. Histology and microanatomy of vertebrae, ribs, haemal arch, and humeri and femora of 10 individuals of Stereosternum and two dorsal ribs of 1 individual of Brazilosaurus were studied. All individuals had achieved a body length of 50 cm (equal to 65 % of the maximum known body length) or larger. All sampled bones are highly osteosclerotic due to the reduction of medullary cavities and the filling of medullary regions by endosteal bone. Calcified cartilage occurs – if at all – only locally in small clusters in the medullary regions of midshaft and in higher amounts only in non-midshaft sections of long bones and towards the medio-distal rib shaft, respectively. The primary bone tissue consists of highly organized parallel-fibred tissue and/or lamellar tissue, which is in most samples relatively lightly vascularized or even avascular. If present, vascular canals are mainly longitudinally oriented; some show a radial orientation. Simple vascular canals as well as primary osteons occur. Some of the latter are secondarily altered, i.e. widened. Remodelling of the periosteal cortex is only documented by few scattered erosion cavities and secondary osteons. The tissue is regularly stratified by lines of arrested growth (LAGs), which usually appear as double or multiple rest lines, indicating strong dependence on exogenous and endogenous factors. Because of the inhibition of periosteal remodelling the growth record is complete and no inner cycles are lost. Individuals of Stereosternum show a poor correlation of body size and number of growth marks, which might be the result of developmental plasticity. Brazilosaurus shows a highly organized, avascular lamellar tissue and a high number of regularly deposited rest lines throughout the cortex of the ribs. The medullary region in the ribs of Brazilosaurus is distinctly larger when compared to ribs of Stereosternum. However, strong osteosclerosis is obvious in both taxa, pointing to a high degree of aquatic adaption. Ribs of Stereosternum, Brazilosaurus, and Mesosaurus are clearly distinguishable from each other by the distribution of the periosteal and endosteal territory. Furthermore, Brazilosaurus differs in its growth pattern (i.e. spacing of rest lines) when compared to Stereosternum and Mesosaurus.

The early Permian mesosaurs were the first amniotes to re-invade aquatic environments. One of their most controversial and puzzling features is their distinctive caudal anatomy, which has been suggested as a mechanism to facilitate caudal autotomy. Several researchers have described putative fracture planes in mesosaur caudal vertebrae -unossified regions in the middle of caudal vertebral centrathat in many extant squamates allow the tail to separate and the animal to escape predation. However, the reports of fracture planes in mesosaurs have never been closely investigated beyond preliminary descriptions, which has prompted scepticism. Here, using numerous vertebral series, histology, and X-ray computed tomography, we provide a detailed account of fracture planes in all three species of mesosaurs. Given the importance of the tail for propulsion in many other aquatic reptiles, the identification of fracture planes in mesosaurs has important implications for their aquatic locomotion. Despite mesosaurs apparently having the ability to autotomize their tail, it is unlikely that they actually made use of this behaviour due to a lack of predation pressure and no record of autotomized tails in articulated specimens. We suggest that the presence of fracture planes in mesosaurs is an evolutionary relic and could represent a synapomorphy for an as-yet undetermined terrestrial clade of Palaeozoic amniotes that includes the earliest radiation of secondarily aquatic tetrapods.As the first group of amniotes to return to an aquatic lifestyle, and a key line of evidence for the theory of continental drift 1 , mesosaurs have figured prominently in reconstructions of early amniote evolution. Mesosauridae is currently composed of three monotypic genera (Mesosaurus tenuidens Gervais, 1865 2 , Stereosternum tumidum Cope, 1886 3 , and Brazilosaurus sanpauloensis Shikama and Ozaki, 1966 4 ), all of which are only known from localities that would have been part of an inland Gondwanan sea during the early Permian. Despite debate regarding the exact placement of Mesosauridae among early reptiles, researchers generally agree that they represent one of the most basal reptile clades 5-9 .The anatomy of mesosaurs has also been debated extensively, and one of the more contentious aspects concerns whether or not they had the capacity for caudal autotomy. Caudal autotomy is the ability to drop a part of the tail in order to escape predation, an anti-predator behaviour that is prevalent in several clades of extant lepidosaurs 10,11 . In extant reptiles, caudal autotomy can occur between caudal vertebrae (intervertebral) or along planes of weakness along the caudal centrum that split a vertebra in two (intravertebral). The latter form of autotomy is the most common among extant reptiles and is the only traceable form of autotomy in the fossil record 12,13 . Regeneration of a cartilage cone posterior to the autotomized region may also occur following autotomy, however, several lineages of lepidosaurs do not regenerate their tails 11 .Several rese...