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.     

Paleobiogeographic significance of a new ozawainelloid fusulinid Pseudoacutella partoazari n. sp., from the Asselian (lowermost Permian) of Gaduk (Central Alborz,Iran)

The uppermost Carboniferous–Lower Permian Dorud Group that crops out at the Gaduk section of Central Alborz is over 44 m thick and includes thick sequences of conglomerate, quartzarenite, calcareous sandstone, oncolitic fusulinid limestone, sandy limestone, sandstone and shale. The Toyeh, Emarat and Shah Zeid formations of this Group were dated as latest Gzhelian to Early Sakmarian. A review of the Asselian fusulinids and smaller foraminifers of Iran emphasizes (1) how the foraminiferal assemblages of Alborz and central Iran differ; (2) why non-Pseudofusulina cannot be a nomen substitutum to Pseudofusulina (even if this latter was often misinterpreted in the Tethyan regions). A new smaller fusulinid Pseudoacutella partoazari n. sp. is described from the Emarat Formation (Asselian) that crops out in the Gaduk area. The palaeobiogeographic significance of Pseudoacutella is discussed, because this genus, which was cosmopolitan from Late Bashkirian to Moscovian, became rare in the Upper Pennsylvanian of the USA and the Carnic Alps, and then, very rare in Lower Permian of Texas, Arizona and Iran.     

Late Ordovician brachiopods from Peru and their palaeobiogeographical relationships

A Sandbian brachiopod association from the Calapuja Formation, in the Peruvian Altiplano, north‐west of Lake Titicaca, has allowed a re‐examination of the palaeobiogeographical relationships between Gondwana and Avalonia during the Late Ordovician, when the palaeocontinents are considered to be already very distant from one another. The brachiopod fauna includes the new species Onnizetina calapujensis sp. nov., Horderleyella chacaltanai sp. nov., Drabovinella minuscula sp. nov. and Tasmanella curtiseptata sp. nov., as well as Caeroplecia sp., Dinorthis cf. flabellulum and Tunariorthis cardocanalis. In addition, Colaptomena expansa expansa and Heterorthis retrorsistria, known from the British Burrellian Stage of the Caradoc Series (late Sandbian) in Wales and the Welsh Borderlands, have also been identified. The brachiopod collection is the most diverse known from a single locality in the whole Central Andean Basin. Within it, forms with clear Gondwanan links occur, such as the new species of Onnizetina, Drabovinella and Horderleyella, and typical representatives of the Avalonian faunas, such as the Welsh Colaptomena expansa expansa and Heterorthis retrorsistria. The brachiopod species exchange between the Proto‐Andean margin of Gondwana and Avalonia, now believed to be possible during the late Sandbian, allows a reconsideration of the global taxonomic affinities of both regions. With this in mind, detrended correspondence analysis (DCA) and cluster analysis have been applied to an updated rhynchonelliformean brachiopod matrix consisting of presence/absence data. The scatter plot resulting from the DCA allows a vivid visualization of the grouping and geographical trends of the South American localities with respect to Avalonia–Baltica and the Mediterranean margin of Gondwana during the Sandbian. Our results agree with previous palaeogeographical reconstructions, depicting Avalonia very close to Baltica and already distant from Gondwana. As a few brachiopod species, with low dispersal potential, would have been able to migrate between those distant palaeocontinents, the existence of intermediate islands in the Rheic Ocean, permitting the transit by island hopping of eurythermal species, must be considered.     

Pliensbachian gastropods from Venetian Southern Alps (Italy) and their palaeobiogeographical significance

Abstract: The Pliensbachian gastropods described by De Toni in 1912, coming from an isolated boulder at the foot of Mt Vedana (eastern margin of Trento Platform, Venetian Southern Alps, Italy) are revised. The fauna consists of 13 species representing nine families and eight superfamilies. Despite the low number of species, the assemblage represents the most diverse Early Jurassic gastropod fauna known for the Venetian Southern Alps. The boulder yielding the material was thought to derive from the upper part of the Early Jurassic Calcari Grigi Group, a carbonate platform unit extensively cropping out in the Mt Vedana area. The sedimentological analysis indicates a prevalently bioclastic wackstone‐floatstone, reflecting a lime‐muddy deposit undergone to an early consolidation. This and the high content of ammonoids, which is unusual for the Calcari Grigi Group, are typical aspects of a condensed pelagic sediment, presumably a fissure filling at the top of the carbonate platform succession. Palaeobiogeographical comparisons show that the fauna is composed of species occurring exclusively in pelagic limestones of the western Tethys. By contrast, it shows no relationships with the coeval faunas of the adjacent Trento platform and of the other western Tethyan carbonate platforms. These lines of evidence and the facies analysis would testify to the Pliensbachian drowning of the eastern margin of the Trento platform. In the wider context of the palaeobiogeographical history of Early Jurassic western Tethyan gastropods, the species from Vedana belong to a faunal stock which is typical for pelagic, mainly postdrowning sediments. Thus, appearance and diffusion in space and time of this stock were probably regulated by the direction, rate and pattern of the Neotethyan rifting. A new subgenus, Proarcirsa (Schafbergia) subgen. nov., and three new species, namely Ataphrus (Ataphrus) cordevolensis sp. nov., Guidonia pseudorotula sp. nov. and Proarcirsa (Schafbergia) zirettoensis sp. nov. are erected.     

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.     

Reappraisal of some Upper Devonian (Famennian) spiriferide brachiopods from the Band-e Bayan Domain (Afghanistan)

The upper and uppermost Famennian succession (Upper Devonian) of the Band-e Bayan Domain (also known as the Axial Zone), a Cimmerian terrane of western Afghanistan (northern margin of Gondwana), yielded diverse brachiopod faunas that are dominated by the spiriferides. Among the latter, six species belonging to four genera, previously erected by D. Brice in the seventies and also known from Iran, are reappraised on the basis of their type material and re-illustrated. The genera Enchondrospirifer and Eobrachythyris are transferred to the subfamilies Cyrtospiriferinae (Cyrtospiriferoidea) and Prospirinae (Spiriferoidea), respectively, whereas the ornament and micro-ornament of both Afghan species of Dichospirifer are documented. The genus Ghorispirifer nov. gen. (Cyrtiopsinae) is erected with Cyrtiopsis graciosa chakhaensis Brice as type species, and includes a second species, namely Cyrtiopsis lapparenti Brice. The new genus is characterized notably by the strong development of ventral apical callosities. Upper Devonian (Frasnian?) brachiopods from the Band-e Bayan Domain described by Maillieux in the first half of the 20th century are revised and photographically illustrated for the first time. The brachiopod generic diversity recognized in Afghanistan (Band-e Bayan Domain, Central Mountains) prior to the Hangenberg Crisis is very high in some sections of the Band-e Bayan Domain (Rhuk pass section, and to a lesser extent at Robat-e-Pai), as also observed in contemporaneous successions of South China and contrary to what is reported in Western Europe (Franco-Belgian Basin for instance).     

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.     

Early Permian brachiopod faunas from Peninsular India: Their Gondwanan relationships

The Early Permian (Late Asselian? to Aktastinian?) brachiopod faunas of Peninsular India are revised in terms of current taxonomy. Genera such as Semilingula, Arctitreta, Etherilosia, Strophalosia, Aulosteges, Bandoproductus, Cyrtella, Neospirifer, Crassispirifer, Tomiopsis and Gilledia confirm the Gondwanan aspect of the faunas and a close relationship, at the generic level, to the Early Permian brachiopod faunas of Western Australia. Peninsular Indian Early Permian brachiopod faunas belong to a complex of Gondwanan and peri‐Gondwanan faunas from Oman and the Pamirs in the West to Australasia in the east. This distribution implies relative freedom of migration for the faunas along the northern margin of Gondwana during the Early Permian.     

Evaluating neurophylogenetic patterns in the larval nervous systems of brachiopods and their evolutionary significance to other bilaterian phyla

Recent structural analyses of invertebrate nervous systems have supported hypotheses stating that specific developmental and cytological aspects of larval and adult brains are conserved among bilaterian animals. Opposing views argue that structural similarities in larval nervous systems may be the result of convergent evolution and that the developmental diversity of adult brains is more indicative of several independent origins. Here, I use various cytological probes, confocal microscopy, and reconstruction techniques to investigate the cellular diversity within the larval nervous systems of Glottidia pyramidata and Terebratalia transversa (Brachiopoda). Neuronal cell types are compared among the rhynchonelliform, linguliform, and craniiform brachiopods as well as the phoronids. Although the respective larval types of the previously mentioned systematic groups clearly diverge in the neuroarchitecture of their larval apical organs (and nervous systems in general), a ground plan is proposed based on shared, centrally‐located, peptidergic neuronal cell types that can be compared with similar cell types in other lophotrochozoan phyla (bryozoans and spiralians). Assessing hierarchal levels of homology within and among the nervous systems of morphologically disparate phyla is challenging in that many phyla share early developmental signals that induce the specification of the neural ectoderm, clouding our ability to discern divergent larval and juvenile brain structure. Solving these problems will require a combined effort involving both traditional and more recent cytological techniques with a diversity of molecular probes that will better map the neuronal complexity of diverse invertebrate nervous systems. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Early Jurassic (latest Toarcian) brachiopods from the northeastern margin of the Western Tethys (Central Iran) and their paleobiogeographical significance

Brachiopod fauna from central Iran, recorded in the upper part of the Shemshak Group and attributed to the upper Toarcian (Pseudoradiosa-Aalensis zones), are reported for the first time in Iran. The assemblage recognized includes six different taxa: Homoeorhynchia sepahanensis nov. sp., formally described in this paper, Globirhynchia subobsoleta, Pseudogibbirhynchia sp., Tetrarhynchiidae sp. indet., Monsardithyris? aff. haresfieldensis, and Zeilleria cf. leckenbyi. Analysis of faunal affinities with other paleobiogeographical regions shows a free connection of the central Iranian brachiopod fauna and wide areas of the northern shelf margin of the Tethys Ocean. This is due to an apparent disruption of bioprovinciality inferred for the late Toarcian-earliest Aalenian, congruent with a connection through the northern seaway across the peri-Laurasian epicontinental platforms.     

Two new late Carboniferous Neuropteris species (Medullosales) from Saarland, Germany and their palaeobiogeographical significance

Two new species ( Neuropteris terminiscus and N. kneuperi ) are described from the Bolsovian ('Westphalian C') of Saarland. N. terminiscus had previously been misidentified as Paripteris pseudograndinioides , but the pinnae are clearly imparipinnate and thus cannot belong to that morphogenus. N. kneuperi had previously been misidentified as Neuropteris obliqua and N. heterophylla , both which appear to be absent from Saar-Lorraine. This is considered further evidence of the distinctive nature of the Westphalian vegetation of the Saar-Lorraine Basin, especially among the seed-plants.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 139 , 193–205.     

Early Devonian (Lochkovian) trilobites and brachiopods from the west Balkhash Region,Kazakhstan and their significance for regional terrane dynamics

A moderately diverse Early Devonian (Lochkovian) trilobite and brachiopod association has been recovered from the lower part of the volcano-sedimentary Shakshagaily Formation, exposed on the western side of Lake Balkhash in central Kazakhstan. Its discovery demonstrates the presence of Lower Devonian marine deposits in the region. The trilobite assemblage includes representatives of six genera: the dalmanitid Kasachstania alperovichi nov. sp. is the dominant taxon, with other components being Ananaspis?, Leonaspis?, Maurotarion, Trimerus (Edgillia), and Warburgella. Associated brachiopods include species of Glossoleptaena, Leptaena, Protolepstostrophia, Pseudostrophochonetes, and Resserella. Whilst a significant proportion of these taxa show distinct links with the contemporaneous trilobite and brachiopod faunas of the north Balkhash Region, the occurrence of Trimerus (Edgillia) aff. kinglakensis Gill, 1949 suggests a possible link with the Lochkovian trilobite fauna of the Australasian sector of Gondwana. The west Balkhash Region preserves an extensive record of late Silurian to Early Devonian island arc volcanism, slope-rise sedimentation and olistostrome formation, which has invariably been ignored in models favouring early amalgamation of the Kazakhstanian terranes into a single continent by the end of the Ordovician.     

Phylogenomic analyses of lophophorates (brachiopods, phoronids and bryozoans) confirm the Lophotrochozoa concept

Based on embryological and morphological evidence, Lophophorata was long considered to be the sister or paraphyletic stem group of Deuterostomia. By contrast, molecular data have consistently indicated that the three lophophorate lineages, Ectoprocta, Brachiopoda and Phoronida, are more closely related to trochozoans (annelids, molluscs and related groups) than to deuterostomes. For this reason, the lophophorate groups and Trochozoa were united to Lophotrochozoa. However, the relationships of the lophophorate lineages within Lophotrochozoa are still largely unresolved. Maximum-likelihood and Bayesian analyses were performed based on a dataset comprising 11,445 amino acid positions derived from 79 ribosomal proteins of 39 metazoan taxa including new sequences obtained from a brachiopod and a phoronid. These analyses show that the three lophophorate lineages are affiliated with trochozoan rather than deuterostome phyla. All hypotheses claiming that they are more closely related to Deuterostomia than to Protostomia can be rejected by topology testing. Monophyly of lophophorates was not recovered but that of Bryozoa including Ectoprocta and Entoprocta and monophyly of Brachiozoa including Brachiopoda and Phoronida were strongly supported. Alternative hypotheses that are refuted include (i) Brachiozoa as the sister group of Mollusca, (ii) ectoprocts as sister to all other Lophotrochozoa including Platyzoa, and (iii) ectoprocts as sister or to all other protostomes except chaetognaths.     

Eocene Pachynolophinae (Perissodactyla,Palaeotheriidae) from China,and their palaeobiogeographical implications

The Eocene perissodactyl family Palaeotheriidae has traditionally been considered to be a nearly endemic European group within Equoidea, but a few palaeotheres have been reported from Asia. Here, I reanalyse a maxilla containing M1–3 from the Lunan Basin, Yunnan Province, China. This element was initially assigned to a new tapiromorph species, Lophialetes yunnanensis, but is here placed in a new genus Lophiohippus within Pachynolophinae based mainly on the absence of mesostyles, the strongly oblique metalophs, the strong development of lophodonty, parastyles overlapping metastyles of preceding teeth and situated mesial to the paracone, and the fact that M3 is longer than wide and has a large and buccally deflected metastyle. Lophiohippus differs from European Anchilophus and Paranchilophus in that the parastyles are situated mesial or even slightly lingual, rather than mesiobuccal, to the paracones, and M3 is markedly relatively larger than M1. I further reanalyse Qianohippus magicus from the Shinao Basin of Guizhou Province, China, in which the complete dentition is known. Qianohippus is characterized by a molariform P2 and non‐molariform P3–4; a relatively high degree of lophodonty; the absence of mesostyles; an angular bending in the protoloph on P3‐M3 and the metaloph on M1–3 at the paraconule and metaconule, respectively; and weakly developed ‘metastylid’ on the lower cheek teeth. A cladistic analysis supports a close relationship between Lophiohippus yunnanensis and Paranchilophus, and suggests that Qianohippus is closely related to some derived pachynolophs. The appearance of the pachynolophins Lophiohippus and Qianohippus in China supports the existence of a biogeographical connection between Europe and Asia in the Middle‐Late Eocene, and the dispersal route was probably along the Tethyan microcontinents in the south.     

Paleogeographic significance of Early Permian crinoids and blastoids from Oman

Early Permian crinoids and blastoids from Oman show relationship with Late Paleozoic Tethyan faunas of Timor and Western Australia and support an Early Permian age for part of the Basleo fauna of Timor. The camerate crinoidPlatycrinites omanensis n. sp. and the blastoidsTimoroblastus andDeltoblastus are reported for the first time from Sakmarian strata of northeastern Oman, doubling the known Permian echinoderms from Oman. The blastoids suggest an offshore lower energy shelf environment of deposition.      

New Early Jurassic brachiopods from the Western Tethys (Eastern Subbetic, Spain) and their systematic and paleobiogeographic affinities

Two new brachiopod genera (Alebusirhynchia and Praesphaeroidothyris), five new species (Alebusirhynchia jorali, Salgirella? goicoecheai, Praesphaeroidothyris cisnerosi, Lychnothyris lancisi, and Cincta peiroi) and a new combination (Securina plicata) have been recorded and established after the internal and external morphological study of populations attributed to Lower Jurassic carbonate successions of the South-Iberian Paleomargin (SE Spain). The analysis made enables us to detail the diagnostic characters of little-known genera such as Lychnothyris or Securina and to clarify the systematics of the group from new supraspecific definitions, establishing a new, strongly multicostate rhynchonellid genus (Alebusirhynchia) and a new smooth terebratulid genus (Praesphaeroidothyris). The palaeobiogeographic signature of these taxa has been analysed, revealing remarkable Mediterranean affinities. Thus, the previously restricted geographical distribution of these faunas can be extended to the westernmost Tethys areas, where the maximum degree of endemism is established for the brachiopod groups in the Subbetic domain around the Sinemurian-Pliensbachian transition.     

New and poorly known acrotretid brachiopods (Class Lingulata) from the Cedaria-Crepicephalus zone (late Middle Cambrian) of the Great Basin, USA

Seven new late Middle Cambrian (Cedaria-Crepicephalus zone) acrotretid species are described from the Great Basin, USA. The species belong to five genera of which Lensotreta is new. The following new species are described: Anabolotreta? glabra, Anabolotreta mogota, Canthylotreta crista, Dactylotreta elegantula, Lensotreta perplexa, Opisthotreta transversa, and Opisthotreta indistincta. The shell structure of Canthylotreta is described for the first time based on new material of the type species Canthylotreta marjumensis and of the new species C. crista. The shell of Canthylotreta includes two types of shell fabric, camerate and columnar. Moreover, parts of the shell show transitional textures, supporting the view that the camerate shell structure may has originated from a columnar fabric. Picnotreta lophocracenta Robson and Pratt, 2001, recently described from western Newfoundland, is also described from the new occurrence in the Great Basin.     

Oldest known Dicoelosia and Epitomyonia,deep water brachiopods from the Beiguoshan Formation (Middle Katian,Upper Ordovician), Shaanxi,north China

Abstract: A diverse brachiopod fauna from a relatively deep water carbonate facies of the Upper Ordovician Beiguoshan Formation (uppermost Caradoc – lower Ashgill, middle Katian) is characterized by small shells and contains the oldest known Dicoelosia and Epitomyonia, two diagnostic taxa of deep water brachiopod palaeocommunities during the Late Ordovician and Silurian. Three new species are recognized: Dicoelosia cordiformis sp. nov., Dicoelosia perbrevis sp. nov. and Epitomyonia fui sp. nov. These pioneer forms of the family Dicoelosiidae show a relatively high degree of morphological plasticity. The shells of Dicoelosia from the Beiguoshan Formation range from the typical slender‐lobed form with a concavoconvex profile to the strongly equibiconvex, fat‐lobed morphotype that was not known previously until the late Silurian. The Beiguoshan dicoelosiids point to an important attribute of the deep water brachiopods: small generalists with high morphological plasticity, which make them ideal candidates as progenitors for the evolution of shallow water brachiopod faunas in shelf and platform depositional environments.     

Permophricodothyris Pavlova, 1965 (Brachiopoda, Spiriferida) from the Permian of South China: its morphology, biostratigraphy and distribution

A study of both silicified and nonsilicified specimens of Permian reticularioid brachiopods from South China suggests thatPermophricodothyris, a genus previously rarely reported from China, is actually very common and abundant in the Middle and especially Upper Permian of South China. This study also clarifies, for the first time, that many of the reticularioid brachiopod species previously described asSquamularia in fact belong toPermophricodothyris. The new data presented in this paper also allows a critical evaluation ofPermophricodothyris in relation to its closest allies:Phricodothyris, Squamularia, Bullarina andNeophricodothyris. The revision reveals that a total of 18Permophricodothyris species are present in the Middle and Upper Permian of South China, with only one species,P. squamularioides, having survived the Permian-Triassic mass extinction. Two species,P. grandis (Chao) andP. guangxiensis Han, Zhou & Wang, are redescribed here, providing critical new information on the morphology and taxonomy of these species.     

<Emphasis Type="Italic">Mongousia</Emphasis>: A new productid genus (Brachiopoda) from the Permian of northeastern Mongolia

A new linoproductid genus is described from the provincial Terrakea arguta Zone (lower part of the Upper Kazanian Substage) of the key section of the Tsagan-Temete Horizon (Kazanian Stage-Lower Tatarian Substage) of northeastern Mongolia. The type species of the new genus was found in the middle layers of the Lower Uldza Subformation, which contains numerous diverse productids from the genus Terrakea.