Basal Therapsid Research Articles

Computed Tomography elucidates ontogeny within the basal therapsid clade Biarmosuchia

Biarmosuchia is a clade of basal therapsids that includes forms possessing plesiomorphic ‘pelycosaurian’ cranial characters as well as the highly derived Burnetiamorpha which are characterised by cranial pachyostosis and a variety of cranial bosses. Potential ontogenetic variation in these structures has been suggested based on growth series of other therapsids with pachyostosed crania, which complicates burnetiamorph taxonomic distinction and thus it is essential to better understand cranial ontogeny of the Burnetiamorpha. Here, three new juvenile biarmosuchian skulls from the late Permian of South Africa are described using X-ray micro computed tomography (CT). We found that juvenile biarmosuchians are distinguished from adults by their relatively large orbits, open cranial sutures, and incomplete ossification of the braincase and bony labyrinth. Also, they manifest multiple centres of ossification within the parietal and preparietal bones. CT examination reveals that the holotype of Lemurosaurus pricei (BP/1/816), previously alleged to be a juvenile, shows no evidence of juvenility and is thus probably an adult. This suggests that the larger skull NMQR 1702, previously considered to be an adult L. pricei, may represent a new taxon. This study provides, for the first time, a list of characters by which to recognise juvenile biarmosuchians.

Biostratigraphy of the Eodicynodon Assemblage Zone (Beaufort Group, Karoo Supergroup), South Africa

Abstract The middle Permian Eodicynodon Assemblage Zone is the lowermost biozone of the Beaufort Group (Adelaide Subgroup, Karoo Supergroup) and occurs in the southwestern part of the main Karoo Basin. It is host to a diverse assemblage of basal therapsid genera of which Eodicynodon is the most abundant. The biozone reaches a maximum thickness of 1 100 m in the Prince Albert Road area and thins to the east and west. The biozone corresponds to the Combrinkskraal and Grootfontein members of the Abrahamskraal Formation, directly overlies the Waterford Formation of the Ecca Group, and records the earliest middle Permian terrestrial environments of Gondwana. Rocks of the biozone were deposited along the southern shoreline of the Karoo Basin in a subaerial delta plain environment as part of large-scale fan systems draining to the north and northeast within a second-order highstand systems tract.

Oxygen isotopes and ecological inferences of Permian (Guadalupian) tetrapods from the main Karoo Basin of South Africa

The Abrahamskraal Formation of South Africa has a diverse middle Permian (Guadalupian) fossil tetrapod fauna comprising largely basal therapsids and pareiasaurian parareptiles. Knowledge on the paleoecology of these animals is limited because inferences of many ecological traits in mid-Permian tetrapods are based on deductions from morphology. While it is widely accepted that the rhinesuchid amphibians were aquatic, the ecological lifestyle of pareiasaurs and dinocephalian therapsids remains uncertain. To investigate the ecological roles of different middle Permian tetrapod taxa we studied stable oxygen isotope compositions of phosphate from teeth and bones as a proxy for water dependence. Our results reveal similar δ18Op values for pareiasaurs, the dinocephalian Anteosaurus, and therocephalians, with a range of values similar to those of extant terrestrial species. The consistent δ18Op values for pareiasaurs strengthen our interpretation of a terrestrial lifestyle for these parareptiles, but the lower Anteosaurus δ18Op values might question the interpretation and a larger sample size may result in a more conclusive outcome. These results conclude the debate on the ecology of middle Permian pareiasaurs. A terrestrial lifestyle is applicable only to the middle Permian species and a different lifestyle is possible for pareiasaurs species that lived after the end Capitanian extinction.

Re-evaluation of an early sphenacodontian synapsid from the Lower Permian of England

ABSTRACTThe holotypic isolated maxilla of the early sphenacodontianHaptodus grandisfrom the Cisuralian of England is revisited. A unique character combination includes haptodontine-grade traits like less specialised teeth and a high number of precanines, but at the same time a shortened lacrimal that is separated from the naris, which is strongly diagnostic of Sphenacodontoidea. As the specimen cannot be assigned to any known taxon, the new taxonHypselohaptodus grandisgen. nov. is proposed. Comparison with other sphenacodontians reveals a mosaic distribution of maxillary features, most significantly regarding the precanine region. Preliminary character histories precludeHypselohaptodusfrom Sphenacodontidae, but suggest a haptodontine-grade or basal therapsid position. The latter hypothesis is substantiated by an ecological model of episodic wet phases in an overall trend of aridification throughout the Permian, to explain the rareness of non-sphenacodontid sphenacodontians in the fossil record. Also from the early Permian of England, an isolated dentary has previously been assigned toOphiacodon, but can be shown to be either a sphenacodontian, possibly affiliated withHypselohaptodus, or a robust,Stereorhachis-like ophiacodontid. The absence ofOphiacodonin the intramontaneous Permian basis of Europe is explained by a narrow environmental tolerance range requiring limnic connection with lowland basins.

The postcranial skeleton of the basal tapinocephalid dinocephalian Tapinocaninus pamelae (Synapsida: Therapsida) from the South African Karoo Supergroup

Dinocephalians form an important component of the Guadalupian basal therapsid faunas of Pangaea. Most research undertaken on this clade has focused on the skull while postcranial research has lagged, largely because of the rarity of sufficiently complete specimens. The discovery of an almost complete skeleton of the basal tapinocephalid dinocephalian Tapinocaninus from the rocks of the lowermost Beaufort Group of South Africa for the first time provides an accurate vertebral count for a dinocephalian as well as morphological information on the appendicular skeleton. The long bones of Tapinocaninus pamelae are autapomorphic in several features within the dinocephalians, and the results enable discussion of some more general aspects of the appendicular skeleton of basal synapsids. Combined, the new data enable a new reconstruction of the posture of tapinocephalids. Although demonstrating several apomorphic characters, the skeleton retains pleisomorphic anatomical features previously known only in pelycosaur-grade synapsids, especially in the axial skeleton. The discovery greatly advances understanding of the postcranial morphology of tapinocephalid dinocephalians and will provide input on the enigmatic phylogenetic relationships of early therapsids.

A new species of burnetiid (Therapsida, Burnetiamorpha) from the early Wuchiapingian of South Africa and implications for the evolutionary ecology of the family Burnetiidae

AbstractBurnetiidae is a family of basal therapsids that is known from sequences of late Permian (Lopingian) age from southern and eastern Africa and European Russia. Recent discoveries of related genera within the broader clade Burnetiamorpha have added to our understanding of morphological variation in the group but have eroded the list of characters defining the family Burnetiidae. We describe a new burnetiid taxon, Leucocephalus wewersi gen. et sp. nov., and argue that Burnetiidae can be defined by, among other characters, the presence of two bosses on the ventrolateral surface of suborbital bar and zygomatic arch, high skull angulation between the orbits, and a median frontal crest that becomes wider and lower posteriorly. The new specimen was found in the early Wuchiapingian Tropidostoma Assemblage Zone of the Main Karoo Basin and, along with previous discoveries, indicates that the family reached its greatest diversity and abundance in the early Wuchiapingian. Diversity declined into the later Wuchiapingian and Changhsingian. Although the clade Burnetiamorpha, including the family Burnetiidae, contains at least 11 genera, each of these is exceptionally rare, with most represented by only one specimen. This could be attributed to a genuine ecological characteristic or may be the result of biogeographical factors, particularly if the Main Karoo Basin was on the periphery of their range.

The mystery of a missing bone: revealing the orbitosphenoid in basal Epicynodontia (Cynodontia, Therapsida) through computed tomography.

The basal non-mammaliaform cynodonts from the late Permian (Lopingian) and Early Triassic are a major source of information for the understanding of the evolutionary origin of mammals. Detailed knowledge of their anatomy is critical for understanding the phylogenetic transition toward mammalness and the paleobiological reconstruction of mammalian precursors. Using micro-computed tomography (μCT), we describe the internal morphology of the interorbital region that includes the rarely fossilized orbitosphenoid elements in four basal cynodonts. These paired bones, which are positioned relatively dorsally in the skull, contribute to the wall of the anterior part of the braincase and form the floor for the olfactory lobes. Unlike procynosuchids and the more basal therapsids in which the orbitosphenoids are well developed, dense, and bear a ventral keel, the basal epicynodonts Cynosaurus, Galesaurus, and Thrinaxodon display cancellous, reduced, and loosely articulated orbitosphenoids, a condition shared with many eucynodonts. The hemi-cylindrical orbitosphenoid from which the mammalian condition is derived re-evolved convergently in traversodontid and some probainognathian cynodonts.

Cranial Bosses of Choerosaurus dejageri (Therapsida, Therocephalia): Earliest Evidence of Cranial Display Structures in Eutheriodonts.

Choerosaurus dejageri, a non-mammalian eutheriodont therapsid from the South African late Permian (~259 Ma), has conspicuous hemispheric cranial bosses on the maxilla and the mandible. These bosses, the earliest of this nature in a eutheriodont, potentially make C. dejageri a key species for understanding the evolutionary origins of sexually selective behaviours (intraspecific competition, ritualized sexual and intimidation displays) associated with cranial outgrowths at the root of the clade that eventually led to extant mammals. Comparison with the tapinocephalid dinocephalian Moschops capensis, a therapsid in which head butting is strongly supported, shows that the delicate structure of the cranial bosses and the gracile structure of the skull of Choerosaurus would be more suitable for display and low energy combat than vigorous head butting. Thus, despite the fact that Choerosaurus is represented by only one skull (which makes it impossible to address the question of sexual dimorphism), its cranial bosses are better interpreted as structures involved in intraspecific selection, i.e. low-energy fighting or display. Display structures, such as enlarged canines and cranial bosses, are widespread among basal therapsid clades and are also present in the putative basal therapsid Tetraceratops insignis. This suggests that sexual selection may have played a more important role in the distant origin and evolution of mammals earlier than previously thought. Sexual selection may explain the subsequent independent evolution of cranial outgrowths and pachyostosis in different therapsid lineages (Biarmosuchia, Dinocephalia, Gorgonopsia and Dicynodontia).

Palaeoneurological clues to the evolution of defining mammalian soft tissue traits.

A rich fossil record chronicles the distant origins of mammals, but the evolution of defining soft tissue characters of extant mammals, such as mammary glands and hairs is difficult to interpret because soft tissue does not readily fossilize. As many soft tissue features are derived from dermic structures, their evolution is linked to that of the nervous syutem, and palaeoneurology offers opportunities to find bony correlates of these soft tissue features. Here, a CT scan study of 29 fossil skulls shows that non-mammaliaform Prozostrodontia display a retracted, fully ossified, and non-ramified infraorbital canal for the infraorbital nerve, unlike more basal therapsids. The presence of a true infraorbital canal in Prozostrodontia suggests that a motile rhinarium and maxillary vibrissae were present. Also the complete ossification of the parietal fontanelle (resulting in the loss of the parietal foramen) and the development of the cerebellum in Probainognathia may be pleiotropically linked to the appearance of mammary glands and having body hair coverage since these traits are all controlled by the same homeogene, Msx2, in mice. These suggest that defining soft tissue characters of mammals were already present in their forerunners some 240 to 246 mya.

Physiological implications of the abnormal absence of the parietal foramen in a late Permian cynodont (Therapsida).

The third eye (pineal eye), an organ responsible for regulating exposure to sunlight in extant ectotherms, is located in an opening on the dorsal surface of the skull, the parietal foramen. The parietal foramen is absent in extant mammals but often observed in basal therapsids, the stem-group to true mammals. Here, we report the absence of the parietal foramen in a specimen of Cynosaurus suppostus, a Late Permian cynodont from South Africa (SA). Comparison with Procynosuchus delaharpeae, a contemporaneous non-mammalian cynodont from SA, demonstrates that the absence of this foramen is an abnormal condition for such a basal species. Because seasonality was marked during the Late Permian in SA, it is proposed that the third eye was functionally redundant in Cynosaurus, possibly due to the acquisition of better thermoregulation or the evolution of specialized cells in the lateral eyes to compensate for the role of the third eye.

Lende chiweta, a new therapsid from Malawi, and its influence on burnetiamorph phylogeny and biogeography

ABSTRACTThe Chiweta Beds of Malawi have yielded a diverse late Permian fossil tetrapod fauna that correlates with that of the Cistecephalus Assemblage Zone of the South African Karoo Supergroup. Amongst the fossil therapsids from the Chiweta Beds is the well-preserved skull and lower jaw of a burnetiamorph, a group of biarmosuchians with numerous bosses and swellings on the skull. This specimen was reported in a preliminary paper in 2005 as the first burnetiamorph described outside of South Africa and Russia. Reanalysis of the morphology and phylogeny of this specimen places Lende chiweta, gen. et sp. nov., as the sister taxon to the clade formed by Proburnetia (Paraburnetia (Pachydectes, Bullacephalus, Burnetia, Niuksenitia)). The greatest diversity of this basal therapsid group is from South Africa, with six of nine described genera and a stratigraphic range that extends from the middle Permian Tapinocephalus Assemblage Zone to the upper Permian Dicynodon Assemblage Zone. Bearing in mind the constraints that govern fossil preservation, current data suggest that what is now southern Africa may have been the area of origin for burnetiamorphs. Under this premise, what is now central Africa represented a corridor that allowed migration of representatives of the group between the southern and northern portions of Pangea during the late Permian.http://zoobank.org/urn:lsid:zoobank.org:pub:BB094736-24AE-4B98-A33CBD7B7668DC2FSUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVPCitation for this article: Kruger, A., B. S. Rubidge, F. Abdala, E. Gomani Chindebvu, and L. L. Jacobs. 2015. Lende chiweta, a new therapsid from Malawi, and its influence on burnetiamorph phylogeny and biogeography. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2015.1008698.

Rise of dinosaurs reveals major body-size transitions are driven by passive processes of trait evolution

A major macroevolutionary question concerns how long-term patterns of body-size evolution are underpinned by smaller scale processes along lineages. One outstanding long-term transition is the replacement of basal therapsids (stem-group mammals) by archosauromorphs, including dinosaurs, as the dominant large-bodied terrestrial fauna during the Triassic (approx. 252-201 million years ago). This landmark event preceded more than 150 million years of archosauromorph dominance. We analyse a new body-size dataset of more than 400 therapsid and archosauromorph species spanning the Late Permian-Middle Jurassic. Maximum-likelihood analyses indicate that Cope's rule (an active within-lineage trend of body-size increase) is extremely rare, despite conspicuous patterns of body-size turnover, and contrary to proposals that Cope's rule is central to vertebrate evolution. Instead, passive processes predominate in taxonomically and ecomorphologically more inclusive clades, with stasis common in less inclusive clades. Body-size limits are clade-dependent, suggesting intrinsic, biological factors are more important than the external environment. This clade-dependence is exemplified by maximum size of Middle-early Late Triassic archosauromorph predators exceeding that of contemporary herbivores, breaking a widely-accepted 'rule' that herbivore maximum size greatly exceeds carnivore maximum size. Archosauromorph and dinosaur dominance occurred via opportunistic replacement of therapsids following extinction, but were facilitated by higher archosauromorph growth rates.

New Basal Synapsid Supports Laurasian Origin for Therapsids

The distant evolutionary ancestry of mammals is documented by a rich therapsid fossil record. While sphenacodontid synapsids are considered the sister-group of therapsids, the place of origin of therapsids is an enigma, largely because of a long standing morphological and temporal gap (Olson's Gap) in their fossil record. We describe a new large predatory synapsid, Raranimus dashankouensis gen. et sp. nov., from the Middle Permian of Dashankou in China which has a unique combination of therapsid and sphenacodontid features. This specimen is of great significance as it is a basal therapsid which is the sister taxon to all other therapsids. The fact that it was found in association with Early Permian tetrapods (Anakamacops and Belebey) suggests that it is the oldest therapsid and provides the first evidence of therapsid-bearing rocks which cover Olson's Gap. It further supports that therapsids may have had a Laurasian rather than Gondwanan origin.

A new specimen of Biseridens qilianicus indicates its phylogenetic position as the most basal anomodont

A new well-preserved basal therapsid skull from the Xidagou Formation, Middle Permian of China, is identified as Biseridens qilianicus. The following synapomorphies distinguish Biseridens as an anomodont and not an eotitanosuchian as previously described: short snout; dorsally elevated zygomatic arch and septomaxilla lacking elongated posterodorsal process between nasal and maxilla. The presence of a differentiated tooth row; denticles on vomer, palatine and pterygoid; contact between tabular and opisthotic; lateral process of transverse flange of pterygoid free of posterior ramus and absence of mandibular foramen exclude it from other anomodonts. Our cladistic analysis indicates Biseridens to be the most basal anomodont, highlights separate Laurasian and Gondwanan basal anomodont clades and suggests that dicynodonts had their origins in the Gondwanan clade. The co-occurrence of the most basal anomodont (Biseridens) together with the most basal therapsid (Raranimus), basal anteosaurid dinocephalians, bolosaurids and dissorophids suggests that the earliest therapsid faunas are from China.

A new burnetiid (Therapsida: Biarmosuchia) from the Upper Permian of South Africa and its biogeographic implications

ABSTRACT The basal therapsid clade Burnetiamorpha is currently known from six genera of Middle-to-Late Permian age. Bullacephalus, Burnetia, Lemurosaurus, and Lobalopex are from the Beaufort Group of South Africa, whereas Niuksenitia and Proburnetia are from Russian deposits. Here we describe a new taxon from the Upper Permian Teekloof Formation of South Africa that is remarkably similar to Proburnetia. Paraburnetia sneeubergensis, gen. and sp. nov., is diagnosed by the following autapomorphies: supratemporal ‘horn’ oriented vertically with bulbous terminal end, supra-orbital boss with well-defined apical crest, and palatine-pterygoid boss elongate. Paraburnetia and Proburnetia share features indicating a sister-taxon relationship, including the presence of a well-developed median nasal crest and tall supraorbital bosses. Along with the close morphological similarity and phylogenetic relationships of several dicynodont taxa from Russia and South Africa, the discovery of Paraburnetia indicates that long-distance, north-south interchange was commonplace during the Late Permian and that burnetiamorphs likely had a Pangean distribution.

A second specimen of Lemurosaurus pricei (Therapsida: Burnetiamorpha)

ABSTRACT A second specimen of the basal therapsid Lemurosaurus pricei preserves aspects of this taxon's cranial anatomy, such as the structure of the temporal region, palate, and lower jaw, that were either unknown or poorly preserved in the holotype. Although typically considered a dinocephalian feature, intermeshing of the incisors is confirmed for Lemurosaurus. Autapomorphies of the genus include a nubbin-like boss at the apex of the lateral temporal fenestra, the presence of very coarse serrations on the posterior margins of the upper postcanine teeth, and the low, ridge-like form of the median frontal crest. The anatomical data provided by the new Lemurosaurus specimen permits the first cladistic appraisal of its phylogenetic position among biarmosuchian therapsids. A cladistic analysis of 10 therapsids, two outgroups, and 38 characters indicates that Lemurosaurus is the sister taxon to a clade including Bullacephalus, Proburnetia, and Burnetia. Taxonomically, this position implies the reclassification of Lemurosaurus as a basal burnetiamorph, rather than as an ictidorhinid. Biogeographically, the position of Lemurosaurus makes it most parsimonious to infer that burnetiamorphs originated in southern Africa, although this hypothesis ignores potential sampling biases.

A new burnetiamorph (Therapsida: Biarmosuchia) from the Lower Beaufort Group of South Africa

The basal clade Burnetiamorpha is known from only two specimens representing two genera, Proburnetia from the Severodvinskian horizon of the Vyatka River Basin in the Kotelnich district of Russia, and Burnetia from the Dicynodon Assemblage Zone of the Beaufort Group of South Africa. Both genera are of Late Tatarian (Late Permian) age. This paper describes the cranial morphology of a new genus of burnetiamorph, Bullacephalus, from the Late Permian Tapinocephalus Assemblage Zone of the Beaufort Group of South Africa. It is known from a relatively complete skull and lower jaw and is the best preserved burnetiamorph yet discovered. Apart from being the oldest member of the clade, Bullacephalus is also morphologically the least derived and provides new evidence on the phylogeny of this poorly understood group of basal therapsids.