Jennifer Botha-Brink

The radiation of cynodonts and the ground plan of mammalian morphological diversity.

Cynodont therapsids diversified extensively after the Permo-Triassic mass extinction event, and gave rise to mammals in the Jurassic. We use an enlarged and revised dataset of discrete skeletal characters to build a new phylogeny for all main cynodont clades from the Late Permian to the Early Jurassic, and we analyse models of morphological diversification in the group. Basal taxa and epicynodonts are paraphyletic relative to eucynodonts, and the latter are divided into cynognathians and probainognathians, with tritylodonts and mammals forming sister groups. Disparity analyses reveal a heterogeneous distribution of cynodonts in a morphospace derived from cladistic characters. Pairwise morphological distances are weakly correlated with phylogenetic distances. Comparisons of disparity by groups and through time are non-significant, especially after the data are rarefied. A disparity peak occurs in the Early/Middle Triassic, after which period the mean disparity fluctuates little. Cynognathians were characterized by high evolutionary rates and high diversity early in their history, whereas probainognathian rates were low. Community structure may have been instrumental in imposing different rates on the two clades.

Osteohistology of the Triassic Archosauromorphs Prolacerta, Proterosuchus, Euparkeria, and Erythrosuchus from the Karoo Basin of South Africa

ABSTRACT The South African non-archosauriform archosauromorph Prolacerta and the archosauriforms Proterosuchus, Erythrosuchus, and Euparkeria were important constituents of the Early to early Middle Triassic Karoo ecosystem following the end-Permian mass extinction. We present new data on the osteohistology of these stem archosaurs and provide insight into their paleobiology. Bone tissues of the Early Triassic Prolacerta contain a poorly defined fibro-lamellar complex, with parallelfibered bone in some regions, whereas the contemporaneous Proterosuchus exhibits rapidly forming uninterrupted fibrolamellar bone early in its ontogeny, which becomes slow forming lamellar-zonal bone with increasing age. The early Middle Triassic Erythrosuchus deposited highly vascularized, uninterrupted fibro-lamellar bone throughout ontogeny, whereas the growth of the contemporaneous Euparkeria was relatively slow and cyclical. When our data are combined with those of previous studies, preliminary results reveal that Early and Middle Triassic non-crown group archosauromorphs generally exhibit faster growth rates than many of those of the Late Triassic. Early rapid growth and rapid attainment of sexual maturity are consistent with life history expectations for taxa living in the unpredictable conditions following the end-Permian mass extinction. Further research with larger sample sizes will be required to determine the nature of the environmental pressures on these basal archosaurs.

Osteohistology of the Triassic archosauromorphsProlacerta,Proterosuchus,Euparkeria, andErythrosuchusfrom the Karoo Basin of South Africa

ABSTRACT The South African non-archosauriform archosauromorph Prolacerta and the archosauriforms Proterosuchus, Erythrosuchus, and Euparkeria were important constituents of the Early to early Middle Triassic Karoo ecosystem following the end-Permian mass extinction. We present new data on the osteohistology of these stem archosaurs and provide insight into their paleobiology. Bone tissues of the Early Triassic Prolacerta contain a poorly defined fibro-lamellar complex, with parallel-fibered bone in some regions, whereas the contemporaneous Proterosuchus exhibits rapidly forming uninterrupted fibro-lamellar bone early in its ontogeny, which becomes slow forming lamellar-zonal bone with increasing age. The early Middle Triassic Erythrosuchus deposited highly vascularized, uninterrupted fibro-lamellar bone throughout ontogeny, whereas the growth of the contemporaneous Euparkeria was relatively slow and cyclical. When our data are combined with those of previous studies, preliminary results reveal that Early and Middle Triassic non-crown group archosauromorphs generally exhibit faster growth rates than many of those of the Late Triassic. Early rapid growth and rapid attainment of sexual maturity are consistent with life history expectations for taxa living in the unpredictable conditions following the end-Permian mass extinction. Further research with larger sample sizes will be required to determine the nature of the environmental pressures on these basal archosaurs.

A mixed-age classed ‘pelycosaur’ aggregation from South Africa: earliest evidence of parental care in amniotes?

Living species of mammals, crocodiles and most species of birds exhibit parental care, but evidence of this behaviour is extremely rare in the fossil record. Here, we present a new specimen of varanopid ‘pelycosaur’ from the Middle Permian of South Africa. The specimen is an aggregation, consisting of five articulated individuals preserved in undisturbed, close, lifelike, dorsal-up, subparallel positions, indicating burial in ‘life position’. Two size classes are represented. One is 50% larger than the others, is well ossified, has fused neurocentral sutures and is distinguished by a coat of dermal ossifications that covers the neck and shoulder regions. We regard this individual to be an adult. The remaining four skeletons are considered to be juveniles as they are approximately the same size, are poorly ossified, have open neurocentral sutures and lack dermal ossifications. Aggregates of juvenile amniotes are usually siblings. Extant analogues of adult and juvenile groupings suggest that the adult is one of the parents, leading us to regard the aggregation as a family group. The Late Middle Permian age of the varanopid family predates the previously known oldest fossil evidence of parental care in terrestrial vertebrates by 140 Myr.

Bone microstructure and the evolution of growth patterns in Permo-Triassic therocephalians (Amniota, Therapsida) of South Africa.

Therocephalians were a speciose clade of nonmammalian therapsids whose ecological diversity and survivorship of the end-Permian mass extinction offer the potential to investigate the evolution of growth patterns across the clade and their underlying influences on post-extinction body size reductions, or ‘Lilliput effects’. We present a phylogenetic survey of limb bone histology and growth patterns in therocephalians from the Middle Permian through Middle Triassic of the Karoo Basin, South Africa. Histologic sections were prepared from 80 limb bones representing 11 genera of therocephalians. Histologic indicators of skeletal growth, including cortical vascularity (%CV) and mean primary osteon diameters (POD), were evaluated in a phylogenetic framework and assessed for correlations with other biologically significant variables (e.g., size and robusticity). Changes in %CV and POD correlated strongly with evolutionary changes in body size (i.e., smaller-bodied descendants tended to have lower %CV than their larger-bodied ancestors across the tree). Bone wall thickness tended to be high in early therocephalians and lower in the gracile-limbed baurioids, but showed no general correlation with cross-sectional area or degree of vascularity (and, thus, growth). Clade-level patterns, however, deviated from previously studied within-lineage patterns. For example, Moschorhinus, one of few therapsid genera to have survived the extinction boundary, demonstrated higher %CV in the Triassic than in the Permian despite its smaller size in the extinction aftermath. Results support a synergistic model of size reductions for Triassic therocephalians, influenced both by within-lineage heterochronic shifts in survivor taxa (as reported in Moschorhinus and the dicynodont Lystrosaurus) and phylogenetically inferred survival of small-bodied taxa that had evolved short growth durations (e.g., baurioids). These findings mirror the multi-causal Lilliput patterns described in marine faunas, but contrast with skeletochronologic studies that suggest slow, prolonged shell secretion over several years in marine benthos. Applications of phylogenetic comparative methods to new histologic data will continue to improve our understanding of the evolutionary dynamics of growth and body size shifts during mass extinctions and recoveries.

A burrow cast with Lystrosaurus skeletal remains from the Lower Triassic of South Africa

Abstract We report on a large burrow cast with skeletal contents from Lower Triassic strata of the Palingkloof Member of the Balfour Formation, which forms the lowermost portion of the Lystrosaurus Assemblage Zone (LAZ) of South Africa. The burrow cast is similar to large burrow casts previously described from the LAZ that were identified as large-scale Scoyenia domichnia. It is the first large burrow cast from the LAZ found to contain diagnostic fossil bone. The burrow cast is a relatively straight, subhorizontal (inclined ∼12°), dorsoventrally compressed tube consisting of an entry ramp and living chamber; the entrance to the burrow is not preserved and there is no evidence that the ramp formed a spiral section. The skeletal material comprises a single, partial, disarticulated skeleton of a juvenile animal that can be assigned with confidence to the dicynodont genus Lystrosaurus. Whereas similar large-diameter burrow casts from strata slightly higher in the LAZ have been attributed to Lystrosaurus, we present an alternative hypothesis that a carnivorous tetrapod constructed the burrow. Our preferred hypothesis is supported by the observation that the interred Lystrosaurus skeleton is too small to be the maker of this particular burrow, by the general observation that carnivorous tetrapods construct relatively straight burrows, and by the partial, disarticulated state of the skeleton, which we interpret as the remains of larded prey. We suggest that akidnognathid theriodonts of the genera Moschorhinus or Olivierosuchus, the most conspicuous large predators of the LAZ, were the constructors of large-diameter, subhorizontal burrows.

Origin of the unique ventilatory apparatus of turtles

The turtle body plan differs markedly from that of other vertebrates and serves as a model system for studying structural and developmental evolution. Incorporation of the ribs into the turtle shell negates the costal movements that effect lung ventilation in other air-breathing amniotes. Instead, turtles have a unique abdominal-muscle-based ventilatory apparatus whose evolutionary origins have remained mysterious. Here we show through broadly comparative anatomical and histological analyses that an early member of the turtle stem lineage has several turtle-specific ventilation characters: rigid ribcage, inferred loss of intercostal muscles and osteological correlates of the primary expiratory muscle. Our results suggest that the ventilation mechanism of turtles evolved through a division of labour between the ribs and muscles of the trunk in which the abdominal muscles took on the primary ventilatory function, whereas the broadened ribs became the primary means of stabilizing the trunk. These changes occurred approximately 50 million years before the evolution of the fully ossified shell.

Anatomy and Relationships of the Middle Permian Varanopid Heleosaurus scholtzi Based on a Social Aggregation from the Karoo Basin of South Africa

ABSTRACT The anatomy of individuals is described from a ‘pelycosaur’ aggregation preserving two age classes, from the Middle Permian of the South African Karoo Basin, that have been referred to Heleosaurus scholtzi. These skeletons allow the recognition of new features for H. scholtzi, including a strong anterior inclination of the occiput, exclusion of the quadratojugal from the temporal fenestra, a maxilla-prefrontal contact, and a maxilla-quadratojugal contact. In addition to some of these features, the absence of an anterodorsal contact between the squamosal and the jugal allows us to assign H. scholtzi to the varanopid clade Mycterosaurinae. Phylogenetic analysis of Varanopidae using an augmented data matrix from the literature positions Heleosaurus as the sister taxon of Elliotsmithia within the varanopid subfamily Mycterosaurinae. Both this sister-group relationship and the monophyly of Mycterosaurinae are weak and collapse with one extra step. Many of the clades discovered in the analysis exhibit the same support level, however, indicating that further studies on varanopids are needed.

Body size and growth patterns in the therocephalian Moschorhinus kitchingi (Therapsida: Eutheriodontia) before and after the end-Permian extinction in South Africa

Abstract The continuous fossil record of therocephalian therapsids (Eutheriodontia) across the Permo-Triassic boundary and their differential survivorship of the end-Permian extinction offer an exceptional deep-time perspective on vertebrate life-history evolution during episodes of large-scale ecological perturbation. To examine potential impacts of the extinction on body size evolution (e.g., “Lilliput” effects) and growth patterns, we investigated cranial sizes and limb bone histology in the therocephalian Moschorhinus kitchingi both before and after the end-Permian extinction, facilitated by analysis of thin-sections of 23 limb bones from an ontogenetic sample of ten individuals across the Permo-Triassic boundary. In general, early subadult Moschorhinus displayed propodial cortices with extensive woven- and parallel-fibered bone (PFB) with dense radial and reticular vascularization and a moderately thickened bone wall with few growth marks. The outer cortex of propodials and epipodials showed a transition to PFB and lamellar bone with longitudinally oriented canals in individuals interpreted as late subadults or adults (>80% largest size). Most elements displayed several (3+) growth marks, though growth marks were more faithfully recorded in the epipodials of Permian individuals. Pearson product-moment correlation tests were performed to examine the relationship between size and robusticity on growth proxies (% cortical vascularity, mean primary osteon diameter), but variation in histomorphology could not be explained by size alone. Variation in body size may be affected by differences in juvenile growth rate and duration, which are highly variable in environmentally stressed extant reptile species. Geologic stage was a more consistent predictor of cortical vascularity. We suggest that Permian and Triassic Moschorhinus exhibited differential rates of early skeletal growth, corroborating the hypothesis that increased environmental variability in the earliest Triassic was associated with rapid growth to a minimum body size requirement and, consequently, shortened developmental times.

Vertebrate Paleontology of Nooitgedacht 68: A Lystrosaurus maccaigi-Rich Permo-Triassic Boundary Locality in South Africa

The farm Nooitgedacht 68 in the Bethulie District of the South African Karoo Basin contains strata that record a complete Permo-Triassic boundary sequence providing important new data regarding the end-Permian extinction event in South Africa. Exploratory collecting has yielded at least 14 vertebrate species, making this locality the second richest Permo-Triassic boundary site in South Africa. Furthermore, fossils include 50 specimens of the otherwise rare Late Permian dicynodont Lystrosaurus maccaigi. As a result, Nooitgedacht 68 is the richest L. maccaigi site known. The excellent preservation, high concentration of L. maccaigi, presence of relatively rare dicynodonts such as Dicynodontoides recurvidens and Dinanomodon gilli, and the large size of many of these Permian individuals makes Nooitgedacht 68 a particularly interesting site for studying the dynamics of the end-Permian extinction in South Africa.