A new basal dicynodont from the Upper Permian of South Africa

The skull of a small anomodont therapsid, from the Tapinocephalus Assemblage Zone (Abrahamskraal Formation, Beaufort Group, Upper Permian) in Northern Cape Province, South Africa, represents a new basal dicynodont and is described in detail. Colobodectes cluveri gen. et sp. nov. is distinguished from other dicynodonts by an anteroposteriorly extensive caniniform process, parietals that were broadly overlapped posterolaterally by posterodorsal processes of the postorbitals, diverging anterior palatal ridges, and a dorsoventrally low foramen magnum. A phylogenetic analysis indicates that Colobodectes is the basalmost member of a dicynodont clade that excludes Eodicynodon . This position is not particularly strong, as two additional steps are needed to make Colobodectes and Eodicynodon oosthuizeni exchange places on the most parsimonious tree. Another discovery of the phylogenetic analysis is that there is little basis for recognizing Eodicynodon oelofseni as the closest relative of E. oosthuizeni . The former species is identified as the sister taxon of a clade that includes the latter and all other dicynodonts.     

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 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.     

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.     

A new fertile Neocalamites from the Upper Permian of Russia and equisetophyte evolution

This paper deals with Neocalamites tubulatus nov. sp., an equisetophyte belonging to the Equisetaceae family. The material studied originates from the Upper Permian deposits of the northern part of European Russia. N. tubulatus nov. sp. is characterized by both sterile and fertile organs belonging to the same parent plant. It has long, flexible stems, rarely branched, with alternating nodes and prolonged ribs on the stem surface corresponding to vascular strands of the stem, and compact lateral strobili, in general aspect similar to the strobili of extant species of Equisetum. N. tubulatus nov. sp. is one of the most ancient representatives of the Equisetaceae family, which shows a distinct morphological pattern of features, among which the lateral position of strobili, characteristic of primitive equisetophytes, is most prominent in this phase of their evolution.     

Zhailimeryx, A New Lophiomerycid Artiodactyl (Mammalia) from the Late Middle Eocene of Central China and the Early Evolution of Ruminants

A new genus and species of primitive selenodont artiodactyl, Zhailimeryx jingweni, occurs in the late middle Eocene Zhaili Member of the Heti Formation, Yuanqu Basin, Shanxi Province, China. A phylogenetic analysis of dental characters suggests that Zhailimeryx is closely related to Lophiomeryx and other lophiomerycid ruminants of Asia and Europe. Zhailimeryx and other Eocene records of Lophiomerycidae from Asia support an Asian origin for this family, followed by dispersal into Europe both before and after the Grande Coupure. Morphological evidence from Zhailimeryx suggests that ruminant artiodactyls had already undergone substantial diversification prior to the late middle Eocene, and it reopens the issue of whether the higher taxa Tragulina and Pecora are valid expressions of ruminant phylogeny.     

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.     

New materials of Cisuralian (early Permian) radiolarians from western Cambodia: Paleobiogeographic implications in the Paleotethys

Radiolarians, including the new species Entactinia pailinensis Ito n. sp., were obtained from the Pailin area, western Cambodia. The radiolarians indicated an age around the Asselian–Sakmarian of the Cisuralian (early Permian). The new species is characterized by an evenly-sized-porous exosphere composed of a polygonal outer porous plate and circular inner porous plate with some radial outer spines. A previous study concluded that Entactinia Foreman rapidly expanded in the early–middle Cisuralian. Our discovery of this new species adds further occurrence data of Entactinia in the early–middle Cisuralian. In addition, another previous study highlighted the uneven distribution of Pseudotormentus De Wever and Caridroit compared with the wider distribution of Quadriremis Nazarov and Ormiston through the Permian, in particular the Guadalupian (middle Permian). Our results combined with those of previous studies indicate that the uneven distribution might extend back to the Sakmarian of the Cisuralian at least.     

Bioerosion in Late Permian Rugosa from reefal blocks (Hawasina Complex,Oman Mountains): implications for reef degradation

Summary Rugose corals are known from allochthonous Late Permian reefal blocks of the A1 Jil and Ba’id Formation (Hawasina Complex), Oman Mountains. In contrast to many Late Permian Rugosa found elsewhere in the Tethys, they occurred in sponge reefs and contributed to reef construction. The waagenophyllid warm water coral fauna is moderately diverse comprising cerioid, thamnasterioid, and fasciculate taxa. In contrast to sponges, chaetetids, and low-growing reefbuilders, the corals secreted diagenetically stable, most probably Mg-calcitic skeletons. Borings in coral skeletons are consequently well preserved providing important data for the interpretation of reef destructive processes. Thin-section analysis revealed three taxa of infaunal borers includingEntobia Bronn 1837, uncertain thallophyte borings, and borings of unknown bioeroders. Macroborers were more important than microborers, because of the dominance of clionid sponges. Good evidence exists also for the occurrence of two types of undetermined grazers which destroyed the coral surfaces. The amount and distribution of bioerosions is variable among different coral taxa. The fasciculate coralPraewentzelella regulare Flügel 1995 was the favorate substrate. Up to 33% of the calices were bored. Dendroid and compound corals were bored subordinately. Bioerosion of these colonies does not exceed 2%. There is good evidence for substrate preference amongst the borers. Major controlling factors affecting borer distribution are believed to be variations of skeletal density and gross morphology. The borer assemblage could not limit reef accretion significantly. Factors controlling boring activity might have been quality of substrate, sedimentation rate, rapid incrustation of substrates, and competition for food with reef constructors including sponges, chaetetids, and rugose corals.     

On aspects of skull form in African apes and orangutans, with implications for hominoid evolution

The study of hominoid phylogeny is currently in a state of controversy and debate due to the discovery of new fossil material and reanalysis of the morphology of extant apes. An important key to the resolution of these debates lies in attaining a fuller understanding of the morphological differences in skull form between the African and Asian great apes. In this paper I have analyzed aspects of facial morphology and internal cranial anatomy in the great apes. Results from this study and previous ones suggest that Pongo is characterized by a marked dorsal deflection of the face relative to the basicranium. Many aspects of circumorbital, midfacial, palatal, and mandibular morphology in Pongo may be related to this airorynchous condition. This hypothesis is supported by Enlow's work on form and pattern in the primate and mammalian skull. The position of the face in known Sivapithecus appears to be similar to that seen in Pongo. Although Pongo may be specialized in its marked degree of airorynchy, it seems likely that an important derived feature linking African apes and hominids is a ventral rotation of the splanchnocranium on the neurocranium. The appearance of marked supraorbital tori and ethmofrontal sinuses are probably correlated developments. Additional implications of this work for debates about hominoid phylogeny are discussed.     

山西柳林孙家沟组锯齿龙类新材料及其分类学意义(英文)

记述了产自山西柳林县薛村镇附近上二叠统孙家沟组的锯齿龙类新材料,包括两个单独的上颌齿和一个保存有牙齿的不完整齿骨。这两颗上颌齿与柳林黄河龙(Huanghesaurus liulinensis)正型标本来自同一地点同一层位且大小相当,由于同一地点没发现过其他的锯齿龙类,推测它们是同种甚至可能是同一个体而且应归入黄河龙;然而在形态特征上这两颗牙齿与矮小三川龙(Sanchuansaurus pygmaeus)正型标本上的牙齿完全相同,可以归入三川龙。新发现的齿骨与矮小三川龙和薛村山西龙(Shansisaurus xuecunensis)正型标本产自同一地点,层位与三川龙相同,低于山西龙的层位,推测很可能也属于三川龙,但是也不排除属于山西龙的可能性;在形态特征上与黄河龙正型标本在齿骨及其下颌齿的形态特征上未见明显差异,可以归入黄河龙。由于柳林黄河龙与薛村山西龙头后骨骼特征也基本一致,认为柳林薛村仅有一个属种的锯齿龙,矮小三川龙和柳林黄河龙应为薛村山西龙的晚出同义名。     

A large anatomically preserved calamitean stem from the Upper Permian of southwest China and its implications for calamitean development and functional anatomy

A large permineralized calamitean stem, Arthropitys yunnanensis Tian et Gu from the Upper Permian of southwest China is reinvestigated and interpreted. The stem has a broad pith and well developed and large carinal canals. Secondary xylem is thick and characterized by wide parenchymatous interfascicular zones that remain constant in width throughout the wood. Striking features of the stem include the abundant leaf traces arranged in two whorls in the cortex with this arrangement previously unrecognized within calamitean stems, and the presence of growth rings in secondary xylem that suggest frequent fluctuations in environmental stress presumably due to variations in water availability. Features of A. yunnanensis infer the stem to be in the epidogenetical phase of calamitean development, and suggest it to be the basal part of a large trunk. Comparisons with biomechanical models for calamitean stems suggest this species had a semi-self supporting habit.     

New latest Permian foraminifers from Laren (Guangxi Province, South China): Palaeobiogeographic implications

Microfacies analyses performed on the latest Permian Wujiaping Formation at Laren (Guangxi Province, South China) show that the bioclastic-rich limestones of Late Permian age contain a rich and well-diversified foraminiferal fauna. This fauna is here revised in order to be compared with time-equivalent levels of southern Iran and southern Turkey. Some new and unexpected phylogenetic trends are highlighted among the biseriamminoids. The new or poorly known genera Retroseptellina, Septoglobivalvulina, Paraglobivalvulinoides, Dagmarita?, Bidagmarita nov. gen., Louisettita, Paradagmaritopsis nov. gen. and Paradagmarita? are concerned. Nevertheless, these newly appeared biseriamminoids are subordinate to abundant Tetrataxis and Climacammina, ultimate survivors of the families Palaeotextulariidae and Tetrataxidae, appeared as old as the Early Carboniferous (“Mississippian”). Algae, miliolids, and nodosarioids are poorly represented. Two genera and four species are here newly described: Globivalvulina curiosa nov. sp., Louisettita ultima nov. sp., Bidagmarita nov. gen., Bidagmarita sinica nov. gen. nov. sp., Paradagmaritopsis nov. gen., Paradagmaritopsis kobayashii nov. gen. nov. sp. The palaeogeographic distribution of these foraminifers is interpreted to be typically of Neo-Tethyan regions, ranging from southern Turkey (Hazro) to South China (Laren) and up to Japan for some species (i.e., Paradagmaritopsis). At Laren, Late Permian strata are generally characterized by Reichelina ex gr. simplex Sheng. Isolated samples of packstones, collected in Tsoteng region (Guangxi Province, South China), contain Sphaerulina sp. together with various smaller foraminifers and numerous representatives of the new species G. curiosa nov. sp. In this study we demonstrate that the regions of Zagros (Iran), Taurus (Turkey), South China and even Japan shared similar foraminiferal assemblages and represented intermittently connected palaeobiogeographic provinces during Late Permian times.     

Thecamoebians from Late Permian Gondwana sediments of peninsular India

The evolutionary history of thecamoebians (testate amoebae) extends back to the Neoproterozoic Era. However, until now, these have had a restricted, discontinuous and modest record across the world. The studied sediment of Raniganj Formation (Godavari Graben), Andhra Pradesh, India has been assigned as Late Permian on the basis of co-occurring age-diagnostic Late Permian palynomorphs. About sixteen thecamoebian species and one taxon incertae sedis have been recorded here in the palynological slides on the basis of shell morphology and morphometry. Out of these, five belong to the family Arcellidae, seven to Centropyxidae, two to Trigonopyxidae, one to Difflugiidae, one to Plagiopyxidae, and one is regarded incertae sedis. The morphometric characteristics of fossil forms resemble their corresponding extant species studied from ecologically diverse fresh water wetlands in India. In general, the ratio of shell diameter and aperture diameter of Late Permian fossil and extant specimens show significant correlation in all the studied species. Except that, the ratio of shell length and breadth is the distinguishing feature between Centropyxis aerophila and C. aerophila ‘sylvatica’, rather than the ratio of shell length and longest diameter of the shell aperture in both fossil and extant forms. The study elucidates the minimal morphological evolution in thecamoebians and their survival during mass extinction periods and stressful environmental conditions over the geological timescale.     

Late Paleozoic stromatolites: new insights from the Lower Permian of Kansas

Carboniferous to Permian marine stromatolites are widely dispersed across the Pangaean margins and embayments and are typified by the ‘Ottonosia-grade stromatolite’ (designated herein). This stromatolite type consists of a well-laminated oncoid or domical stromatoid that developed into branching, laminated columns in the upper reaches. To develop a model for the global pattern, we investigated Lower Permian stromatolites from Kansas (Howe Limestone Member, Red Eagle Limestone). Stromatoids from the Lyon County locality typify the Ottonosia-grade stromatolites. The laminae are sharp throughout the stromatoid and are defined by an increase in cornuspirid foraminfera and algal filaments. The upper zone of the stromatoid is composed of well-laminated branching and brecciated columns (‘pseudo-thrombolitic’). Coeval stromatolites from a new exposure at the Tuttle Creek Dam spillway possess a more massive mesostructure. These stromatolites are composed of a turbinate stromatoid or oncoid base and an overlying domical stromatoid, and are rimmed by smaller meandering columns. Only the basal stromatoid, oncoids, and upper columns are well laminated. In both localities, the microbial-constructing ecosystem is dominated by cornuspirids and calcifying filamentous algae (?Girvanella). The mesostructural differences of the stromatolites are due to different environments of formation. The Tuttle Creek stromatolites formed in a shallow-subtidal to intertidal open marine setting. The coeval Lyon County stromatolites formed in a semi-restricted, marginal marine environment such as a lagoon or supratidal zone. Based on this information and independent sedimentological data, we conclude that lagoonal or supratidal zones were common features in the late Paleozoic intracratonal zones of the Pangaean supercontinent and account for Ottonosia-grade stromatolites occurring in the Laurentian mid-continent, the Zechstein Basin, Japan, Brazil, and Tunisia.     

Permian and Triassic Radiolaria from Northwest Thailand: paleogeographical implications

Well-preserved radiolarians were recovered from seven sections in the Mae Hong Son-Mae Sariang area, northwestern Thailand. 51 species assigned to 34 genera are identified, including 1 new species (Triassospongosphaera erici Feng sp. nov.) and 19 unidentified species. They are divided into the Late Permian, late Ladinian and middle Carnian radiolarian assemblages. Newly identified radiolarian assemblages, together with the published radiolarian biostratigraphic data from this region, indicate that there was a pelagic basin during the Late Paleozoic and Triassic. This basin was joined to the Chiang Dao and Changning-Menglian oceanic basins, and they represent the main oceanic basin of the Paleotethyan Archipelago Ocean. This main oceanic basin was situated in the traditional “Shan-Thai Block”. Therefore, “the Shan-Thai Block” was not a single block during that stage, but composed of the Paleotethyan Ocean and two continental terranes that were affiliated with the Gondwana and Cathaysian domains, respectively.     

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.     

Aragonitic dasycladalean algae from the Upper Permian of Sichuan, China

Exceptionally well-preserved specimens of Mizzia yabei (Karpinsky) were discovered in bituminous packstone intercalations of the early Upper Permian Maokou Formation of Anxian County (Sichuan, China). Staining techniques and x-ray diffraction reveal that the organic encrustation around the central stem and the lateral branches consists of aragonite. Average aragonite contents range between 27% and 40%. The bulk of the remaining 60–73% calcite represents cement. The total Sr-content is 3, 400 ppm, indicating 8,000–10,000 ppm for the aragonite fraction. The organically precipitated aragonite was deposited as closely packed fibres, 10–20 μm long and 0.5–1 μm wide. The fibres are arranged in clinogonal (water-jet) fascicles, which diverge towards the central (spar-filled) stem. This microstructure shows that the mineralization of the mucilage started at the periphery of the extracellular encrustation and progressed toward its interior. Diagenetic alterations of the original aragonite include various stages of breakdown of the fibres into 0.5–1 μm large granules (micritization), minor syntaxial growth of the fibres and local transformation of acicular aragonite into neomorphic calcite. The dasycladaleans from the Permian of Sichuan appear to be by far the oldest representatives of this order with preserved aragonitic mineralogy and microstructure. Comparably well-preserved examples are only known from Eocene and younger deposits.     

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.