A New Basal Lystrosaurid Dicynodont from the Upper Permian of South Africa

A new genus and species of late Permian dicynodont, Kwazulusaurus shakai , is described on the basis of a complete skull from the late Permian Dicynodon Assemblage Zone of the South African Beaufort Group. It is an advanced form which shows characters, such as the shape of the snout and the loss of the ectopterygoid, that link it to the early Triassic genus Lystrosaurus. Kwazulusaurus represents the most basal member of the lystrosaurian lineage. The phylogeny of progressive pristerodontian dicynodonts is discussed. It appears possible that the Kannemeyeriiformes and Lystrosauridae do not form a monophylum, as previously assumed. Instead a sister-group relationship between lystrosaurids and dicynodontids plus kannemeyeriiforms is suggested.     

A new dicynodont (Anomodontia, Eotherapsida) from the Upper Permian of Tatarstan

A new dicynodont genus and species, Idelesaurus tataricus sp. nov. (Cryptodontidae), from the Semin Ovrag locality (Tatarstan, Tetyushinskii District; Upper Permian, Upper Tatarian Substage, Severodvinian Horizon) is described. The skull patterns of the East European and South African Cryptodontidae and Aulocephalodontidae are compared from the morphofunctional point of view.     

A second varanopseid skull from the Upper Permian of South Africa: implications for Late Permian 'pelycosaur' evolution

Late Permian terrestrial faunas of South Africa and Russia are dominated taxonomically and ecologically by therapsid synapsids. On the basis of a single specimen from the Upper Permian of South Africa, the varanopseid Elliotsmithia longiceps is the sole basal synapsid ('pelycosaur') known from Gondwana. Recent fieldwork in the Upper Permian of South Africa has produced a second varanopseid specimen that is referrable to Elliotsmithia . Data from both this specimen and the holotype suggest that Elliotsmithia forms a clade with Mycterosaurus from the Lower Permian of North America and Mesenosaurus from the Upper Permian of Eastern Europe. That postulate is supported by the three most parsimonious trees discovered in a new analysis of varanopseid phylogeny. However, the available data cannot resolve the interrelationships of these three genera. The new phylogenetic results contrast with earlier work identifying Elliotsmithia as the basal member of a clade that includes the North American taxa Aerosaurus , Varanops , and Varanodon . The new trees reduce the stratigraphic debt required by the latter scenario, and the one with the least stratigraphic debt identifies Elliotsmithia and Mesenosaurus as sister taxa. Two new taxa are erected, Mycterosaurinae and Varanodontinae, for the two varanopseid subclades.     

A new galesaurid (Therapsida: Cynodontia) from the Lower Triassic of South Africa

A new galesaurid cynodont, Progalesaurus lootsbergensis gen. et sp. nov., is described on the basis of a well-preserved skull, lower jaw, right scapula and left atlantal neural arch. Autapomorphies of Progalesaurus include postcanine teeth bearing numerous mesial and distal accessory cusps that flank a recurved main cusp, a post-temporal fenestra bordered by the squamosal ventrally and a large external naris. Progalesaurus is similar to Galesaurus in possessing a poorly defined masseteric fossa on the dentary, a strongly recurved main cusp of the postcanine dentition, an incomplete secondary palate and a similar basisphenoid-parasphenoid morphology. A cladistic analysis of ten early cynodont genera resolves a monophyletic Galesauridae encompassing Cynosaurus , Progalesaurus and Galesaurus , although support for this clade is weak. Procynosuchus and Dvinia are placed at the base of Cynodontia whereas Thrinaxodon and Platycraniellus are positioned higher, but outside of Eucynodontia. The holotype and only known specimen of Progalesaurus was collected during systematic prospecting of Permo/Triassic boundary strata at New Lootsberg Pass, Graaff-Reinet District, South Africa. The discovery of Progalesaurus increases the number of valid Early Triassic cynodonts to four and sheds light on the tempo of early cynodont diversification after the end-Permian mass extinction.     

A primitive anomodont therapsid from the base of the Beaufort Group (Upper Permian) of South Africa

Patranomodon nyaphulii , known from a nearly complete skull, lower jaw and partial postcranial skeleton, is morphologically the most primitive anomodont therapsid yet discovered. It is from the Eodicynodon Assemblage Zone, the lowest biozone of the Beaufort Group of South Africa, which has a primitive therapsid fauna comparable with that of the Russian Zone I. Patranaomodon is primitive with respect to other anomodonts in having short palatal exposure of the premaxilla, an unreduced tabular, a slit-like interpterygoidal vacuity, a screw-shaped jaw articulation (which precludes fore-aft sliding of the lower jaw), and only three sacral vertebrae. The poorly-known Galechirus and Galepus from the younger Cistecephalus Assemblage Zone appear to be at a comparably primitive evolutionary grade, and the three genera are tentatively united in the family Galechiridae. The taxon Dromasauria is shown to be paraphyletic and therefore should be discarded.     

A new species of Crossidium (Pottiaceae,Bryophyta) from South Africa

Abstract

A new species of Crossidium (Pottiaceae, Bryophyta), C. karoo, is described for South Africa. It uniquely differs from congeners in tight encapsulation of the photosynthetic filaments by the clasping upper lamina.     

Latest Permian Palaeolithocycliidae (Radiolaria) from South China

The family Palaeolithocycliidae Kozur and Mostler was formerly thought to be ranging from Devonian to Carboniferous. However, a lot of radiolarians assignable to this family were discovered in the late Changhsingian (Latest Permian) of South China. Six species, including Palaeolithocyclia pilata n. sp., Palaeolithocyclia platta n. sp., Paramphibrachium woni n. sp., Klaengspongus spinosus Sashida, Klaengspongus formosus n. sp. and Klaengspongus umbilicatus n. sp., have been identified. In this paper, these species are illustrated and described.     

First reports of Grania (Clitellata: Enchytraeidae) from Africa and South America: molecular phylogeny and descriptions of nine new species

In this article, our knowledge of the geographic distribution of Grania species is expanded by describing seven new species, G rania bekkouchei sp. nov. , G rania brasiliensis sp. nov. , G rania capensis sp. nov. , G rania chilensis sp. nov. , G rania cryptica sp. nov. , G rania hinojosai sp. nov. , and G rania simonae sp. nov. , from poorly investigated regions of the Southern Hemisphere, plus two new species, G rania carolinensis sp. nov. and G rania unitheca sp. nov. , from off the east coast of the USA. An immature achaetous specimen that we call Grania cf. levis was also included. The newly generated data were combined with a previously published data set in order to update the hypothesis of phylogenetic relationships among Grania species. All new species except G . cryptica sp. nov. are supported by both morphological and molecular data. In addition, we find that G . chilensis sp. nov. is structured in at least four distinct populations along the Chilean coast. The species described from South Africa form a monophyletic clade where two are morphologically indistinguishable but diverging in both mitochondrial and nuclear ribosomal genes, and thus we describe them as different species, G . bekkouchei sp. nov. and G . cryptica sp. nov. Among the North Carolinian species, G . unitheca sp. nov. is indicated as a close relative of Grania monospermatheca Erséus & Lasserre, 1976, and G . carolinensis sp. nov. is indicated as a close relative of G. cf. levis. The updated phylogeny is strongly concordant with geographical species distributions, thus supporting a low level of dispersal within this genus, as has previously been hypothesized. © 2015 The Linnean Society of London     

The Upper Permian articulate Phyllotheca australis from South Africa

RAYNER, R. J., 1992. The Upper Permian articulate Phyllotheca australis from South Africa. The taxonomic position of two fossil articulates, which form an important element of the Permian Gondwana flora, is examined in the light of recent collections from South Africa. Vegetative and fertile axes, dispersed and attached sporangia, in-situ spores, isolated and attached whorls of leaves, rhizomes bearing roots and branches, naked axes and a permineralized axis are all assigned to Phyllotheca australis. The large numbers of axes with attached sporangia allow a reinterpretation of the fertile structure and the specific diagnosis is emended. Although it is recognized that the fossils assigned to this taxon may represent a species complex, there exist no clear morphological discontinuities between the material previously assigned to P. australis and P. indica from widely separated Gondwanan localities. Phyllotheca australis was common throughout Gondwana, and is a reliable index fossil of the Upper Permian.     

An Odontogriphid from the Upper Permian of Australia

A problematic organism, Bowengriphus perphlegis gen. et sp. nov., is described based on two specimens from the Late Permian Rangal Coal Measures of eastern central Queensland. It displays a double-looped feeding apparatus bearing small conical elements, considered homologous with that of the supposed lophophorate Odontogriphus omalus Conway Morris, 1976, from the Middle Cambrian Burgess Shale of British Columbia, Canada. Odontogriphids are thus interpreted as a group that survived through the Palaeozoic and made the transition from marine to freshwater environments. Recent proposals that odontogriphids are stem-group chordates are not well established.     

The jaw function and adaptive radiation of the dicynodont mammal-like reptiles of the Karoo basin of South Africa

The structure and functioning of the dicynodont jaw system are described. A pivoting action of the lower jaw around the palate posteromedian to the caniniform processes is as basic to the jaw movement as is the well-known double-convex jaw articulation. The sequence of origin of the characteristic features of the dicynodonts is analysed cladistically; this demonstrates the patterns of association of these characters into functional character-complexes. The structures of the palate and lower jaw, and their functional integration in feeding, are described and illustrated in standard format. Five different lineages of dicynodont can be identified: Eodicynodon; the robertoids (including Diictodon ); the dicynodontoids (including Dicynodon and the majority of the large dicynodont genera of the Permian and Triassic); the endothiodontoids (including Prodicynodon [='Chelydontops'] and Pristerodon ), and the emydopoids (including Cistecephalus, Myosaurus and Kingoria ). Eodicynodon or a similar form could have been ancestral to the other four lineages. The robertoids probably fed upon the stems and rhizomes of equisetaleans, while the varied dicynodontoids probably fed upon the varied glossopterid seed-ferns. The endothiodontoids, too, were probably herbivorous, but many, perhaps all, of the small emydopoids were burrowing and may have been omnivorous. The dicynodonts were probably ectothermal, and the dicynodontoids may have migrated to warmer latitudes in the winter. Only c. 20 genera of Karoo dicynodont are now recognized as valid, and it is suggested that this fauna is now almost completely known. Their distribution in the Karoo biozones is reviewed and correlated with environmental changes. The Permian ancestors of the Triassic dicynodonts, including Lystrosaurus , probably lived on higher, drier ground, and were therefore already adapted to the more fibrous food that spread into the basins as the climate became drier in the Triassic.     

Latest Permian Spumellaria and Entactinaria (Radiolaria) from South China

A diverse and well-preserved latest Permian radiolarian fauna was obtained from bedded siliceous rocks and siliceous mudstones of the Dongpan Section, southwest Guangxi, China. All encountered specimens of the order Spumellaria and a few specimens of the order Entactinaria were selected for the taxonomic study of this paper. Thirty-three species, 15 of which are new species, were identified. They belong to 15 genera, three of which (Paracopicyntra, Copiconulus, and Yujingella) are new. The most interesting result of our study is that eight typical Middle Triassic spumellarian genera (Paroertlispongus Kozur and Mostler, Tetrapaurinella Kozur and Mostler, Paurinella Kozur and Mostler, Lithelius Haeckel, Archaeospongoprunum Pessagno, Tamonella Dumitrica, Kozur and Mostler, Bistarkum Yeh, and Tetraspongodiscus Kozur and Mostler) were discovered in the upper Changhsingian strata studied herein. Therefore, the most important and characteristic spumellarian families of the Middle Triassic were already present in the Late Permian of South China.     

A new Early Permian reptile and its significance in early diapsid evolution

The initial stages of evolution of Diapsida (the large clade that includes not only snakes, lizards, crocodiles and birds, but also dinosaurs and numerous other extinct taxa) is clouded by an exceedingly poor Palaeozoic fossil record. Previous studies had indicated a 38 Myr gap between the first appearance of the oldest diapsid clade (Araeoscelidia), ca 304 million years ago (Ma), and that of its sister group in the Middle Permian (ca 266 Ma). Two new reptile skulls from the Richards Spur locality, Lower Permian of Oklahoma, represent a new diapsid reptile: Orovenator mayorum n. gen. et sp. A phylogenetic analysis identifies O. mayorum as the oldest and most basal member of the araeoscelidian sister group. As Richards Spur has recently been dated to 289 Ma, the new diapsid neatly spans the above gap by appearing 15 Myr after the origin of Diapsida. The presence of O. mayorum at Richards Spur, which records a diverse upland fauna, suggests that initial stages in the evolution of non-araeoscelidian diapsids may have been tied to upland environments. This hypothesis is consonant with the overall scant record for non-araeoscelidian diapsids during the Permian Period, when the well-known terrestrial vertebrate communities are preserved almost exclusively in lowland deltaic, flood plain and lacustrine sedimentary rocks.     

Early Permian bioevent in the fusulinacean fauna of South China

Analysis of a large database of the stratigraphic distribution of fusulinacean Foraminifera reveals an Early Permian event of significant decline of species diversity in South China. Data from Late Carboniferous to Early Permian sections without apparent unconformity in southwest China were evaluated to determine if the apparent pattern of species disappearance was caused by bias in fossil preservation associated with Early Permian sea-level changes. Statistical analysis suggests that the Early Permian event started in the Late Sakmarian with a significant drop of species diversity in the Robustoschwagerina ziyunensis Zone and continued through the Pamirina darvasica Zone of the Artinskian and into the Brevaxina dyhrenfurthi Zone of the Early Kungarian, resulted in a total loss of about 40% species diversity in the fusulinacean fauna. The Early Permian event is the most extensive bioevent in the history of fusulinacean Foraminifera at the species level although it is less significant at the generic level. Because a similar faunal change has been found among the fusulinacean assemblages in North America and in various regions of Tethys, this event may represent a major faunal turn-over in response to the Early Permian changes in sea level and could be of a global nature. Previous recognition of this event was hampered by Early Permian unconformities in North America and other regions of Tethys.     

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 basal sauropodiform from South Africa and the phylogenetic relationships of basal sauropodomorphs

We present a new medium‐sized basal sauropodomorph, S efapanosaurus zastronensis gen. et sp. nov. , from the Upper Triassic?Lower Jurassic Elliot Formation of South Africa. It is represented by parts of the postcranial skeleton of at least four individuals, including: cervical, dorsal, sacral and caudal vertebrae, most of the forelimb, and part of the hindlimb. Sefapanosaurus bears several autapomorphies of the astragalus, and referred material also shows autapomorphic features. The inclusion of Sefapanosaurus in a phylogenetic analysis places it within the group of sauropodomorphs more closely related to sauropods than to Massospondylus (i.e. Sauropodiformes), increasing the currently known diversity of the so‐called ‘transitional forms’ leading to Sauropoda. Character optimization revealed the presence of several features that are common for taxa placed within the transitional branches basal to Sauropoda. Sefapanosaurus, together with other transitional sauropodomorphs reported during the last decade, highlights the importance of Gondwanan taxa for understanding the palaeobiodiversity, global distribution, and macroevolutionary changes in the group related to the rise of sauropods. © 2015 The Linnean Society of London     

Spiral trace fossils from the Permian Ecca Group of Zululand

The presence of spiral trace fossils is reported for the first time from six localities in strata of the Vryheid Formation of the Ecca Group (Lower Permian) in South Africa. These localities are all in the northeast part of the main Karoo basin of sedimentation. The fossils arc assigned to the ichnospecies Spirodesmos archimedeus , representing the spiral trail or burrow of a deposit-feeding organism. In contrast to other reports of spiral trace fossils, the Spirodesmos traces described here were formed in a shallow-water environment. This is established on both sedimentary and ichnofacies evidence. Associated trace fossils include Skolithos, Corophioides and Siphonichnus , all of which arc members of the Skolithos ichnofacies of Scilacher. The occurrence of Spirodesmos in this ichnofacies suggests that these strata were deposited in a marine basin.     

A new, enigmatic family for new genus and species of Polyneoptera from the Upper Permian of Russia

Alexarasniidaefam. n. and Alexarasnia rossicagen. et sp. n. are described from the Upper Permian of European Russia. Systematic position of this enigmatic family within the infraclass Polyneoptera is unclear.     

A new species of Discinites (Noeggerathiales) from the Upper Permian of Weibei Coalfield, North China

A new species of Discinites is described from the Late Permian Upper Shihhotse Formation of the Weibei Coalfield, Shaanxi Province, China. D. hanchengensis sp. nov. has decurrent and disk-like sporophylls with deep-toothed apices, elliptical sporangia and rectangular epidermal cells. The spores are generally of the Calamospora type. The megaspores and microspores are 300–380 and 45–90 μm in diameter, respectively. The microspores are similar in size to those of all other species of Discinites that contain Calamospora, but the megaspores are remarkably smaller. The new species represents the first record of Discinites with Calamospora from China and from the Cathaysian flora. Finally, we summarize the associated foliages of Discinites in the Cathaysian flora and find that they are all markedly different from those found in the Euramerican flora.     

Gryllolattids (Insecta, Grylloblattida) from the Upper Permian of South Africa

A Late Permian fauna of grylloblattid insects (Insecta; Grylloblattida) from the KwaZulu-Natal Province (South Africa) is considered. New taxa, Liomopterites ulterior sp. nov. and L. meridionalis sp. nov. (family Liomopteridae); Normandienoptera serotina gen. et sp. nov. and Afrogrylloblattus disputabilis gen. et sp. nov. (Skaliciidae), are described. Mioloptera stuckenbergi Riek, 1973 and Mioloptoides andrei Riek, 1976 of the family Liomopteridae are redescribed.