The Permian fusulinoidean faunas of the Sibumasu and Baoshan blocks: their implications for the paleogeographic and paleoclimatologic reconstruction of the Cimmerian Continent

Permian fusulinoidean faunas occur in mainly four stratigraphic levels in the Baoshan Block of West Yunnan and the Sibumasu Block of Southeast Asia, which constituted part of the eastern Cimmerian Continent. The oldest fauna, from the upper part of the Dingjiazhai Formation in the Baoshan Block, consists of Pseudofusulina, Eoparafusulina, and a new boultoniid genus, and is assignable to the Yakhtashian (=Artinskian). The second one, which occurs in the basal part of the Ratburi Limestone and its equivalent strata in the Sibumasu Block, is represented by Monodiexodina, and is probably referable to the Bolorian (=Kungurian). The third fauna, composed of Eopolydiexodina, Rugososchwagerina, Yangchienia, Chusenella, Jinzhangia, and several other genera, is dated to the Murgabian (=Wordian), and occurs in the lower part of the Shazipo and Daaozi formations in the Baoshan Block and the main part of the Ratburi Limestone in the Sibumasu Block. The youngest fauna of probably Dzhulfian (=Wuchiapingian) age is found in the upper part of the Ratburi Limestone, and contains Nanlingella, Reichelina, Codonofusiella?, and a few staffellid genera. A smaller foraminiferal genus, Shanita, found from the upper part of the Ratburi Limestone and the upper part of the Shazipo Formation is also an important element of the foraminiferal assemblage near the Midian-Dzhulfian (=Capitanian-Wuchiapingian) boundary in the Baoshan and Sibumasu blocks.In the eastern Cimmerian Continent, low generic diversity throughout the Permian and the paucity of Tethys-characterizing neoschwagerinid and verbeekinid genera during Middle Permian time are two remarkable features of the Permian fusulinoidean faunas. In the Cimmerian Continent, the generic diversity of Permian fusulinoidean faunas in space and time gradually increases from the Early Permian to late Middle Permian as well as from the eastern Cimmerian areas to western ones. The temporal increase of the generic diversity can be explained by the northward drift of the Cimmerian Continent during Permian time. In contrast, the lower generic diversity of the eastern Cimmerian Permian fusulinoidean faunas against western ones is possibly due to an oblique arrangement of the continent to paleolatitude. Thus, the western Cimmerian Continent was more proximal to the tropical Tethyan domain than its eastern part. In addition, the Middle Permian Cimmerian paleobiogeographic region is likely to be subdivided into two subregions, the western Tethyan Cimmerian and the eastern Gondwanan Cimmerian, based on the distribution pattern of verbeekinid and neoschwagerinid fusulinoideans and overall generic diversity. The scarce occurrence or total absence of these essentially Tethys-indicating fusulinoideans in the Baoshan and Sibumasu blocks suggests that the eastern Cimmerian Continent was still far from the equatoro-tropical Cathaysian domain and was probably in a warm temperate or subtropical zone until the end of the Permian. The eastern Cimmerian areas finally migrated into a tropical zone by the Late Triassic judging from well-developed Carnian sponge-coral buildups in the Chaiburi Formation in the Sibumasu Block.     

Changhsingian (Late Permian) foraminifers from the topmost part of the Xiala Formation in the Tsochen area,central Lhasa Block,Tibet and their geological implications

A foraminiferal fauna consisting of 15 genera and 19 species is documented for the first time from the topmost part of the Xiala Formation in the Aduogabu area in Tsochen County, central Lhasa Block. This foraminiferal fauna is dominated by Colaniella, indicating a broadly Changhsingian age. The presence of Colaniella, Reichelina and absence of the typical tropical fusuline genus Palaeofusulina in the fauna indicate that the Lhasa Block has not merged into equatorial regions during the Changhsingian. More importantly, the widespread Lopingian (Late Permian) marine carbonates with warm-water faunas and the stable platform carbonate sequence through the Guadalupian and Lopingian in the Lhasa Block present an obvious contrast against the contemporaneous sequences in the South Qiangtang Block and Gondwana margin. It suggests that both the Bangong-Nujiang Ocean and the Neotethys Ocean would have opened by the Guadalupian and Lopingian.     

A Wuchiapingian (Late Permian) brachiopod fauna from an exotic block in the Indus–Tsangpo suture zone, southern Tibet, and its palaeobiogeographical and tectonic implications

A brachiopod fauna including 19 species of 17 genera from an exotic block in the Indus–Tsangpo suture zone in southern Tibet is described and illustrated. The brachiopod fauna is dominated by Martinia elegans and two new taxa: Jinomarginifera lhazeensis gen. et sp. nov. and Zhejiangospirifer giganteus sp. nov. The fauna is closely comparable with those from the middle and upper parts of the Wargal Formation and the Chhidru Formation in the Salt Range of Pakistan, the Chitichun Limestone in southern Tibet, and the Basleo area of West Timor, and these correlations suggest a Wuchiapingian age. The fauna exhibits substantial links with both peri–Gondwanan and Cathaysian faunas, which may imply that it is a seamount biota originally located in the southern margin of the Neotethys during the Late Permian, and was later (in the early Cenozoic) displaced and became sandwiched into younger marine deposits in the collision process between India and Eurasia.     

A new species of the marattialean fern Scolecopteris (Zenker) Millay from the uppermost Permian of Guizhou Province, south-western China

Several isolated marattialean synangia and sporangia are reported from coal balls collected from Coal Seam No.1 (C605) in the uppermost Permian Wangjiazhai Formation in Guizhou Province, south-western China. The synangia are radially symmetrical with diameters between 0.8 and 1.2 mm and are 1.7 mm long, consisting of 3–4 elongate sporangia that are fused basally, free distally and possess a pointed apex. The outer-facing sporangial wall is 4–5 cells thick and conspicuously differentiated. Spores are trilete, have a granular ornamentation and are nearly round equatorially with a diameter of 55–60 µm. Comparisons with other anatomically preserved Palaeozoic marattialean synangia from the Euramerican and Cathaysian floras permit their assignment to the genus of Scolecopteris (Zenker) Millay. In this species the thick, outer-facing sporangial walls and large trilete spores are features consistent with those of the Oliveri Group within Scolecopteris , a group that has previously been considered primitive within this genus. Distinctions from all other previously recognized species within the Oliveri Group lead to the creation of a new species, S. guizhouensis sp. nov. This species is the youngest of the reported species of Scolecopteris recognized from the Euramerican and Cathaysian floras, and provides important evidence on the organization of marattialean ferns from the Upper Permian strata of south China.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 279–288.     

Middle-late Asselian (Early Permian) fusulinid fauna from the post-Variscan cover in NW Anatolia (Turkey): Biostratigraphy and geological implications

The earliest autochthonous cover of the Variscan basement of the Sakarya Composite Terrane (SCT) in NW Anatolia is represented by basal conglomerates and limestones. The microfacies types of the limestones in ascending order are: (1) bioclastic grainstone/packstone, (2) fusulinid grainstone/packstone, (3) smaller foraminiferal grainstone/packstone, (4) Anthracoporella (dasycladale) grainstone/packstone, and (5) wackestones. Twenty-three species assignable to 15 genera of fusulinids were recovered from the studied materials of the Kadirler section; Quasifusulinaguvenci nov. sp. and Pseudoschwagerinabeedeimagna nov. subsp. are created. Rugosofusulinids, sphaeroschwagerinids, pseudoschwagerinids, occidentoschwagerinids, pseudochusenellids, quasifusulinids, rugosochusenellids and paraschwagerinids are the main faunal elements of the succession, which shows two distinct faunal intervals. Eoschubertella, Schubertella, Biwaella?, Rugosofusulinastabilis group, and Pseudochusenella correspond to the first interval at the base; the second interval is characterized by the species of Sphaeroschwagerina, Pseudoschwagerina,Occidentoschwagerina, the Rugosofusulinalatispiralis group, and diverse quasifusulinids. A biostratigraphic correlation shows that the Kadirler section in the SCT in NW Anatolia shares many common species with Central Asia in the East but especially with the Carnic Alps and Karavanke Mountains in the West. The new data suggest that the close faunal relationship in the Late Carboniferous between eastern Alps, Ural Mountains, NW Turkey and Central Asia also continued during the Asselian.     

Functional aspects of the postcranial anatomy of the Permian dicynodont Diictodon and their ecological implications

The postcranial skeleton of the Permian dicynodont Diictodon is described, major skeletal muscles restored and functional aspects of the skeleton analyzed. The humerus was caudolaterally oriented and articulated in a near sagittal plane. This reduced the transverse component of thrust during locomotion. Throughout the stride, the femur, which is relatively longer than the humerus, was held in a parasagittal position. Diictodon probably had a hindlimb gait similar to the high walk of crocodilians. Comparisons between several dicynodont genera, including Diictodon, reveal considerable variation in the limb orientations, articulations, posture and gait among these taxa. Expansion of the preacetabular iliac process and the overall iliac length with respect to its height among these dicynodonts may be correlated with the increase in the number of the sacral vertebrae. Diictodon does not show the cranial and forelimb modifications suitable for digging as seen in Cistecephalus, Kawingasaurus and extant burrowers such as a broad skull, rounded occiput for considerable neck musculature, a robust humerus and a prominent olecranon process. However, a cylindrical body, short limbs with well–developed limb elevators and depressors and a long wide manus with long, blunt claws probably helped in digging. In addition, the hindlimb with well–developed retractors and short, blunt claws participated in soil removal. The caudolaterally oriented humerus and a laterally flexible vertebral column facilitated movement in the narrow confines of the burrows. Forelimb articulation and morphology indicate that its method of digging was probably rotation–thrust, where humeral excursion and not forearm extension played a dominant role. The close associations of articulated paired skeletons of adult individuals suggest that Diictodon was gregarious. A large number of unconnected burrow–casts in a small area of about 500 m², indicates that although the animals lived close together, they did not share a single large communal structure.     

Jurassic radiolarites in a Tethyan continental margin (Subbetic, southern Spain): palaeobathymetric and biostratigraphic considerations

A region of the pelagic Subbetic basin within the Southern Iberian Continental Margin is studied in lithostratigraphical and biostratigraphical detail. Jurassic radiolarites (Jarropa Radiolarite Formation, Bathonian–Oxfordian) interbedded with shallow-water marine limestones have been recognized. Underlying the radiolarites (Camarena Formation, Bajocian) are oolitic limestones showing shallowing-upward cycles with karstic surfaces on the top, corresponding to deposition on an isolated carbonate platform on volcanic edifices. The Milanos Formation (upper Kimmeridgian–Tithonian), overlying the radiolarites, contains calciclastic strata with hummocky cross-stratification, which indicate outer carbonate ramp deposition. In the Jarropa Radiolarite Formation some calcisiltite strata with hummocky cross-stratification have been found. The bathymetry of the Subbetic Jurassic pelagic sediments, including the radiolarites, is considered as moderate or shallow in depth. We suggest that the pelagic character of the Jurassic sediments in this margin and their equivalents in other Alpine domains is a consequence of distance from the continent (beyond the pericontinental platform) but not necessarily of depositional depth.     

An upper Kungurian/lower Guadalupian (Permian) brachiopod fauna from the South Qiangtang Block in Tibet and its palaeobiogeographical implications

A brachiopod fauna including 15 species belonging to 14 genera is described from a thin carbonate succession of the Lugu Formation at the Anmu section in the northern part of the South Qiangtang Block, which is about 15 km south to the Longmu Co-Shuanghu Suture. The brachiopod fauna is late Kungurian or early Guadalupian in age in terms of the presence of the brachiopod species Vediproductus punctatiformis, “Cryptospirifer” omeishanensis, and Paraplicatifera regularis. This is also generally consistent with the age indicated by the fusulines (e.g., Cancellina, Chusenella, Monodiexodina, Nankinella, Neofusulinella, Pseudofusulina, and Sphaerulina) from this formation at the nearby Mari and Duoma A (DMA) sections. Palaeobiogeographically, the brachiopod fauna exhibits a typical Cathaysian affinity and all the species are reported commonly from the equivalents in South China and Cimmerian blocks, which strongly suggest that the South Qiangtang Block had drifted to a position in the warm-water or temperate regime close to South China.     

Late Pennsylvanian to Middle Permian revised algal and foraminiferan biostratigraphy and palaeobiogeography of the Lycian Nappes (SW Turkey): Palaeogeographic implications

Shallow-marine limestones associated to a Palaeotethyan seamount in the Teke Dere unit of the Tavas Nappe (Lycian Nappes, SW Turkey) are essentially latest Moscovian-Kasimovian in age. The wide range of microfauna and -flora of the series show biogeographic affinities comparable to those from the northern Palaeotethyan borders (especially to assemblages from the Carnic Alps, Urals, Donbass and Darvaz). These biogeographic affinities seem to persist until the end of the Early Permian (Artinskian). The Middle Permian fauna is represented by the typical warm, tropical assemblages known at the same time in the Palaeotethys (NW Caucasus, Darvaz, south China, Primorie and Japan), and in the Neotethys (Transcaucasia, central Iran, southern Afghanistan and Sibumasu). The new Kasimovian algae and incertae sedis Novantiellopsis elliottii n. gen. n. sp., Uvanellopsis fluegelii n. gen. n. sp., Tubiphytes rauzerae n. sp. and Asselodiscus davydovi n. sp. are described.     

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.     

The ontogeny of the Late Permian terebratulids (Brachiopoda) Gruntelasma Smirnova, Grigorjevaelasma Smirnova, and Campbellelasma Smirnova and their taxonomic position

A specific type of ontogenetic change combining elements of dielasmoid and angustothyridid stages of brachidium development is established for the Late Permian genera Gruntelasma Smirnova, 2004, Grigorjevaelasma Smirnova, 2004, and Campbellelasma Smirnova, 2004 from the Russian Platform. On this basis, these genera are assigned to the superfamily Compositelasmatoidea Smirnova, family Compositelasmatidae.     

Palaeoginkgoxylon zhoui, a new ginkgophyte wood from the Guadalupian (Permian) of China and its evolutionary implications

A new taxon of ginkgophyte affinity Palaeoginkgoxylon zhoui gen. nov. et sp. nov. is described from the Guadalupian Lower Shihhotse Formation of the Hulstai coalfield, Inner Mongolia Autonomous Region (Nei Mongol), northern China, on the basis of the anatomical structures of the broad eustele and pycnoxylic secondary xylem. The anatomical structure of the new woody tree trunk resembles both the early gymnosperms of Eristophyton-Pitus types and the modern Ginkgo. Therefore, the new tree trunk is interpreted as representing a transitional stage in the evolution of Ginkgo from early arborescent lignophytes since the Early Carboniferous.     

Rhabdomesid and cystoporid bryozoans from the Permian of Deh-e Mohammad, Shotori Mountains (northeastern Iran)

Seven bryozoan species belonging to the Order Rhabdomesida and Order Cystoporida are described from the Permian deposits exposed near the small town of Deh-e Mohammad, Shotori Mountains (northeastern Iran): Rhabdomeson cf. consimile Bassler, Pamirella nitida Gorjunova, Clausotrypa conferta Bassler, Streblotrypa (Streblotrypa) elegans Sakagami, Streblotrypa (Streblascopora) supernodata nov. sp., Cystodictya sp., and Filiramoporina cf. kretaphilia Fry and Cuffey. The described fauna identifies the age of the Jamal Formation at the locality near Deh-e Mohammad as Lower Permian. It displays palaeobiogeographic connections to the Lower Permian of Pamir (Tajikistan), Indonesia, Thailand and Kansas (North America).     

Cranial morphology and evolution of Permian Dinomorpha (Eotherapsida) of eastern Europe

A Revision of the cranial morphology of Middle and Late Permian East European Dinomorpha (Theromorpha, Eotherapsida) resulted in a clarification of the taxonomic relationships within this group. As the system was constructed, four parameters, i.e., morphological, biomorph, and geographical and stratigraphic ranges, were taken into account in each subordinate taxon. The taxon Theromorpha is ranked as class, since it was primarily formed as a pilidosic group (covered with hair), in contrast to the class Reptilia, a pholidosic group (covered with scales).     

A new species of Dyoros (Brachiopoda; Chonetoidea) from Nevada (United States) and stratigraphic implications for the Pennsylvanian and Permian Antler Overlap assemblage

Newly discovered fossil localities in coarse-grained deposits of the Pennsylvanian and Permian Antler overlap assemblage in the southern Shoshone Range, north-central Nevada have yielded a low-diversity assemblage consisting chiefly of a new species of chonetoidean brachiopod: Dyoros (Lissosia) nevadaensis nov. sp. The subgenus Dyoros (Lissosia), is known from Leonardian and lower Guadalupian strata in North America, mainly in Texas.The coarse-grained lithology of the host strata, their unconformable relation on deformed lower Paleozoic rocks, and the Leonardian and(or) lower Guadalupian age indicated by Dyoros (Lissosia) provide evidence that host strata are younger than strata of the Antler overlap assemblage in nearby areas of the southern Shoshone Range and suggest that an unconformity of local extent may be present within the overlap assemblage. The fossil age ranges and lithologic data suggest that the host strata may be correlative with the Guadalupian Edna Mountain Formation, an unconformity-bounded unit that forms the upper part of the Antler sequence in the Battle Mountain area to the north. This correlation suggests that the unconformity beneath these strata may have regional extent in north-central Nevada. The origin of the inferred regional unconformity is unknown and may have resulted from relative changes of sea level or regional extensional or contractional tectonism in the area of the former Antler highlands, which forms the substrate for the Antler overlap assemblage.     

Forest snail faunas from Transylvania (Romania) and their relationship to the faunas of Central and Northern Europe

Forty‐three forest sites in seven sampling areas in mountainous parts of Transylvania were sampled to obtain inventories of their snail faunas, and to make comparisons between these and the faunas of similar sites further north along the Carpathian chain. Sampling areas ran from close to the Ukrainian border in the north to Retezat in the south‐west. Altogether, 83 species were found, as well as between 19 and 40 species at individual sites. Sites within the same sampling area had more species in common than in among‐area comparisons, although differences between areas did not relate to the distance between them. Such differences appear to be related more to ecological factors than to geography. Faunas of all areas were very similar to those recorded from the Polish and Ukrainian Carpathians; distance from these northern faunas had little effect on similarity. More broadly, the fauna of these forests shows a much greater affinity to those of countries to the north than to those of countries to the south. Despite the survival of forests through the last glacial period of the Pleistocene, and the greater proximity to potential Mediterranean refugia, these faunas appear to be mainly a subset of those found further north. They represent a rather small proportion of the known Romanian fauna, in which there are many endemics restricted to other habitats. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 471–479.     

Guadalupian and Lopingian (Middle and Late Permian) deposits from Mexico and Guatemala, a review with new data

This work describes stage by stage the biostratigraphy of the Middle to Late Permian in Mexico and Guatemala. Roadian deposits are very poorly represented, as a consequence of tectonic movements at the end of the Kungurian/Leonardian stage. In fact Middle and Late Permian deposits are almost completely lacking in South Mexico and the whole Latin America, due to a probable climatic barrier. The main data concern the Las Delicias sequences from Coahuila, North Mexico, and the Mixteca Terrane, South central Mexico, with some precisions on the Wordian-Capitanian from Los Hornos (Puebla) and from Olinalá, Guerrero, respectively with the discoveries of Parafusulina sellardsi and Polydiexodina capitanensis. New data are provided on Capitanian mudmounds from Olinalá. A hypothetical reconstruction of the different terranes of Mexico at the Pangea stage, is finally presented.     

Microornamentation of Lioestheria, Mimoleaia, and Echinolimnadia (conchostraca) from the terminal Permian of Siberia

Previously unknown minute ornamentation patterns of conchostracans are described based on SEM investigation of several collections from the terminal Permian volcanic deposits of the Tunguska Syneclise. Lioestheria (Lioestheriidae) shows ovally elongated cells arranged in rows along the growth bands, Mimoleaia (Leaiidae) has large polygonal cells with double walls, and Echinolimnaia (Echinolimnadiidae fam. nov.) displays a pattern with small polygonal cells bearing small spines along their walls. Diagnoses of genera and species are refined. New data about their distribution are presented.     

Hyloidichnus bifurcatus Gilmore, 1927 and Limnopus heterodactylus (King, 1845) from the Early Permian of Southern Alps (N Italy): A New Equilibrium in the Ichnofauna

Studies on Early Permian tetrapod ichnofauna emphasized the scarcity of forms from Italian sites. A revision work on the entire collections revealed the presence of Hyloidichnus bifurcatus Gilmore, 1927 and Limnopus heterodactylus (King, 1845). The ichnoassociation now lists seven ichnogenera: Amphisauropus, Batrachichnus, Dromopus, Erpetopus, Hyloidichnus, Limnopus, Varanopus. These new data enlarge the ichnoceonosis, adding tracks of medium-size captorhinomorphs (Hyloidichnus) and temnospondyls (Limnopus) to the Italian ichnofauna, previously characterized by scarcity of predators and amphibians. Radiometric ages give a strong age constraint to the ichnoassociation (Early Kungurian), allowing useful correlations to contemporary successions all over the world. The main difference is the absence of Ichniotherium and Dimetropus, and this could have a stratigraphic or paleoenvironmental significance. The fauna is similar in two main basins, Collio and Orobic. It differs solely in the proportions between ichnotaxa, with a predominance of areoscelid traces (Dromopus) in the Collio Basin and of captorhinomorph traces (Erpetopus, Varanopus, Hyloidichnus) in the Orobic Basin. This datum could reflect slightly different environments, seasonal in the Collio Basin (alluvial plain) and more arid in the Orobic Basin (playa-like). The lack of some forms in smaller basins of the Athesian Volcanic Complex is probably due to a bias.