Sheinwoodian (Silurian) conodonts and graptolites from NE Anti‐Atlas,Morocco

Männik, P., Loydell, D.K. & Lubeseder, S. 2010: Sheinwoodian (Silurian) conodonts and graptolites from NE Anti‐Atlas, Morocco. Lethaia, Vol. 44, pp. 410–416. Conodonts and graptolites from a limestone at the base of the Tamaghrout Formation indicate that this is of early Wenlock age. This is the first unequivocal dating of a Sheinwoodian limestone from Morocco. All of the conodonts are known also from several other regions and suggest that there was no major difference in conodont faunas in northern Gondwana, Baltica and Laurentia at this time. Biostratigraphy, conodonts, graptolites, Morocco, Silurian, Sheinwoodian.     

河南内乡晚奥陶世石燕河组牙形石及其地质意义

将河南内乡晚奥陶世石燕河组牙形石自下而上建立了Ｏｕｌｏｄｕｓｕｉｒｉｃｈｉ－Ａｐｈｅｌｏｇｎａｔｈｕｓｓｈｏｓｈｏｎｅｎｓｉｓ带，Ｏｎｏｄｕｓｒｏｂｕｓｔｕｓ带，Ａｐｈｅｌｏｇｎａｔｈｕｓｇｒａｎｄｉｓ带，Ａｐｈｅｌｏｇｎａｔｈｕｓｄｉｖｅｒｇｅｎｓ带和Ａｐｈｅｌｏｇｎａｔｈｕｓｚｕｏｑｕｅｎｓｉｓ带。这个序列与北美中大陆的很相似，两者可详细对比。充分证实了北美中大陆型牙形石动物群以及中国合此类型牙形石的最高奥陶纪地层在华北地台以南的武当地块北缘的存在，并指明了其古生物地理分区和构造意义。     

Allometry in Anisian (Middle Triassic) segminiplanate conodonts and its implications for conodont taxonomy

Conodonts are a clade of chordates and are valuable indicator fossils for biostratigraphy. The segminiplanate (neogondolelliform) conodonts represent a major morphological group ranging from upper Carboniferous to Upper Triassic marine sediments. However, the morphological similarity of segminiplanate P₁ elements generates problems for taxonomy, especially in the Permian and Triassic clades. This paper represents the first study of morphological variation in Triassic segminiplanate conodonts using a geometric morphometric approach. The laminar microstructures observed in conodont cross‐sections indicate that, within our analysed specimens, smaller conodonts with fewer laminae are generally from an earlier ontogenetic stage while larger conodonts with more laminae are from a later stage of ontogeny. Using linear regressions between relative warp scores from both upper and lateral views and conodont length, we demonstrate strongly allometric growth patterns for the species Paragondolella bifurcata Budurov & Stefanov. Our results indicate that the species‐group taxon Pg. praeszaboi bystrickyi (Kovacs et al.) is an early growth stage of Pg. bifurcata and thus synonymous. We suggest that the allometry of conodonts should be considered seriously, especially when there are numerous transitional morphologies between large‐ and small‐sized conodonts. Reconstructing the ontogenetic series and using larger‐sized conodonts within the numerous transitional morphologies in the population of a rock sample for the definition of new species are suggested for future studies.     

Overgrowths of large authigenic apatite crystals on the surface of conodonts from Cantabrian limestones (Spain)

Conodonts from the middle to upper Paleozoic limestones of the Cantabrian zone commonly show apatite overgrowths. A large crystal microtexture observed under the SEM corresponds to local rims of euhedral to subhedral apatite crystals, which were preceded by the neoformation of smaller crystals. Four types of this microtexture (blocky, columnar, fan, and denticular) are described on different areas of the oral surface of conodonts, whereas dissolution features may be present in the basal cavity area. The distribution of these types of microtexture in different areas of conodont morphology suggests a general trend to neocrystallization, where crystal size increases towards the top of the conodont ornamentation and a chemical gradient controls the crystalline growth. This arrangement is widely related to the surface morphology and to the general conodont histology. The large crystal microtexture grows during early diagenesis from near surface to moderate burial and is linked to the known secondary apatite cement present in natural fused clusters of conodonts. The features described here are also compared to microtextures developed on conodonts during low- to medium-grade metamorphic conditions, where phosphate in solution is available.     

A predictive template for the apparatus architecture of the Carboniferous conodont Idioprioniodus

The skeletal architecture of conodonts of the order Prioniodinida Sweet, 1988, is poorly understood. Undisturbed or slightly disturbed natural assemblages are rare. A three-dimensional apparatus plan has been constructed using scale models of the elements of Idioprioniodus , based on late Mississippian and late Pennsylvanian (Carboniferous) assemblages from Montana and Illinois, USA. The interpretation of the architecture is based on symmetry relationships between the various elements. A revised element notation scheme is proposed for the apparatus. □ Conodonts, apparatus, palaeobiology, Prioniodinida , Idioprioniodus.     

依据牙形刺确定的蒙古国塔琳波格德组（Taliin Bogd Formation）的时代

在蒙古南部波尔海尔汗乌尔(Bor hairhan uul)剖面的塔琳波格德组下部的6个样品中发现了牙形刺,通过研究清楚地表明含牙形刺动物群的塔琳波格德组下部的时代是中洛霍考夫阶(早泥盆世)到中艾菲尔阶(中泥盆世).塔琳波格德组的牙形刺动物群更接近于北美同时期的动物群,而与欧洲同期动物群相远.这一动物群可能属于冷水动物群.作者在文中描写了一个新种:Steptotaxis mongolianus sp.nov.Wang.     

BROMALITES FROM THE SOOM SHALE LAGERSTÄTTE (UPPER ORDOVICIAN) OF SOUTH AFRICA: PALAEOECOLOGICAL AND PALAEOBIOLOGICAL IMPLICATIONS

Abstract:  Bromalites from the Soom Shale are allocated to five main categories on the basis of shape, content and internal structure: those containing broken conodont elements; those containing brachiopod fragments; corrugated/spiral forms; coiled coprolites and wrinkled coprolites. It is impossible to allocate specific bromalites to the organisms that formed them, but the occurrence of crushed discinoid valves in several specimens demonstrates that an effective durophagous predator was present in the Soom Shale community. The presence of fragmented elements of conodonts in other specimens provides direct evidence of tiering within the predatory trophic level in the Soom Shale. Conodonts, other agnathan vertebrates, orthoconic cephalopods and eurypterids are all possible contenders for producing most of the bromalites recorded, but there may have been unrecorded large predators in the community.     

The sharpest tools in the box? Quantitative analysis of conodont element functional morphology

Conodonts have been considered the earliest skeletonizing vertebrates and their mineralized feeding apparatus interpreted as having performed a tooth function. However, the absence of jaws in conodonts and the small size of their oropharyngeal musculature limits the force available for fracturing food items, presenting a challenge to this interpretation. We address this issue quantitatively using engineering approaches previously applied to mammalian dentitions. We show that the morphology of conodont food-processing elements was adapted to overcome size limitations through developing dental tools of unparalleled sharpness that maximize applied pressure. Combined with observations of wear, we also show how this morphology was employed, demonstrating how Wurmiella excavata used rotational kinematics similar to other conodonts, suggesting that this occlusal style is typical for the clade. Our work places conodont elements within a broader dental framework, providing a phylogenetically independent system for examining convergence and scaling in dental tools.     

四川盆地下三叠统牙形刺及其油气意义

作者对四川盆地２４条以海相碳酸盐岩地层为主的下三叠统剖面作了系统的研究,在取得了丰富的牙形刺资料的基础,对下三叠统牙形刺进行了区域对比,划分了７个牙形刺带。并对牙形刺色变与油气的关系等进行了研究。编制了ＣＡＩ、古地温等值线图及其与油气分布关系图,并提出了该盆地最有利的油气远景区ＣＡＩ＝２－３;有利的产油气区ＣＡＩ＝２．５－３的新看法。     

Compostional variations and patterns of condont reworking in Late Devonian and early carboniferous calciturbidites (Moravia, Czech Republic)

Summary Compositional variations and grain-size properties of both carbonate constituents and conodonts as an alternative component group were used for interpreting the processes governing the deposition of upper Famennian and middle Tournaisian calciturbidites in Moravia, Czech Republic. Both the composition and grain-size properties of conodont element associations showed to be markedly dependant on facies type of their host sediment. Upper Devonian calciturbidite successions deposited on flanks of wide, Moravian-Silesian carbonate platform are composed mainly of echinoderm-and peloid-rich wacke/packstones and intraclastic float/rudstones (fine-grained calciturbidites, “normal” calciturbidites with Tab Bouma sequences, debris-flow breccias) with abundance of shelf-and shelf margin conodont taxa and epipelagic and “mesopelagic” conodonts. Upper Devonian calciturbidites deposited on slopes of volcanic sea-mounts are composed of echinoderm-and peloid-rich wacke/packstones and float/rudstones with increased proportion of intraclasts and volcanigenic lithoclasts (fine-grained calciturbidites, normal calciturbidites), yeilding abundant conodont associations with higher proportion of “mesopelagic” taxa compared to the platform-flank examples. Middle Tournaisian calciturbidite succession composed of crinoid-, peloid-, intraclast-and lithoclast-rich lime mudstones, wacke/packstones and float/rudstones (normal calciturbidites and debris-flow breccias) yielded conodont element associations rich in shelt-and shelf-margin taxa, “mesopelagic” conodonts and reworked Middle-and Upper Devonian conodonts. In general, the ratio of shelf-and shelf margin conodont taxa to “mesopelagic” taxa is distinctly lower in finegrained calciturbidites than it is in normal calciturbidites and debris-flow breccias. Grain-size properties (mean grain size and sorting) and percentage of fragmented conodont elements, too, are markedly dependant on the facies type: in fine-grained calciturbidites the values of mean grain-size and fragmentation are low and the sorting is good to very good whereas in normal calciturbidites and debris-flow breccias the values of mean grain-size and fragmentation are distinctly higher and the sorting is poorer. The interdependence of facies type and composition and grain-size properties of conodont element associations in gravity-flow deposits is explained as resultant from hydrodynamic sorting during turbidity current flow and final deposition of the bed. Compositional variations observed in our sections may thus be attributed to facies variability (coarsening-and thickening-upward trends) rather than to sea-level fluctuations (highstand shedding of carbonate platforms). On the other hand, significant enrichment in reworked conodont taxa in middle Tournaisian normal calciturbidites compared to scarcity and/or absence of such conodonts in essentially identical facies of upper Famennian age indicate sea-level to be the major control governing such compositional variations, with low relative sea-level stand in middle Tournaisian and high relative sea-level stand in upper Famennian. Thorough analysis of conodont evolution, palaeoecology and taphonomy, with emphasis on understanding the processes of deposition of their host rock, are recommended for any biostratigraphic and biofacies study to be done in carbonate sediments deposited under strong hydrodynamic regimes, such as calciturbidites, temperstites, debris-flow deposits, shelf-edge oolitic sands, tidal-channel facies etc.     

中国二叠系的底界

牙形刺是二叠系生物地层的主导门类,地叠系的三统九阶的界线层型（ＧＳＳＰ）都将用牙形刺定义。我国石炭－二叠系（Ｃ／Ｐ）的分界应采用与国际上一致的标准,即用牙形刺Ｓｔｒｅｐｔｏｇｎａｔｈｏｄｕｓ ｉｓｏｌａｔｕｓ的首次出现定义二叠系的底界。Ｓ．ｉｓｏｌａｔｕｓ在我国华南、华北均有分布,可精确地确定二叠系的底界。华北太原组应归下二叠统Ａｓｓｅｌｉａｎ阶,我国Ｃ／Ｐ界线地层和Ｓｔｒｅｐｔｏｇｎａｔｈｏｄｕ     

Skeletal ontogeny and feeding mechanisms in conodonts

The growth and function of the conodont skeletal apparatus have important implications for early vertebrate relationships and the evolution of vertebrate hard tissues, yet they are poorly understood. Analysis of element length, platform linear dimensions, and platform area in discrete Pa elements of Carboniferous Idiognathodus and Gnathodus bilineatus reveals that the platform increased in size at a rate significantly above that required to maintain geometric similarity. Measurements of P, M and S elements in bedding-plane assemblages of Idiognathodus and G. bilineatus indicate that relative to Pa element length, Pb and S element growth was isometric, whereas M elements grew with negative allometry. There is no evidence to support loss or resorption of S and M elements in later growth stages, or to indicate periodic shedding and replacement of elements. These results are important for understanding apparatus and element Function. The positive allometry of the Pa element platform supports interpretations of a mashing or grinding tooth-like Function for platformed Pa elements. If conodonts were active suspension-feeders, the increasing food requirements of a growing conodont would require the filter array formed by the S and M elements to have grown at a rate significantly above isometry. The lack of positive allometry of S and M elements indicates that conodonts were not suspension-feeders and supports hypotheses that conodonts fed with a raptorial apparatus and teeth. □ Conodonts, vertebrates, skeletal apparatus, ontogeny, allometry, function, suspension-feeding, teeth.     

Preliminary assessment of the use of electron backscatter diffraction (EBSD) in conodonts

Pérez‐Huerta, A., Cusack, M. & Méndez, C.A. 2011: Preliminary assessment of the use of electron backscatter diffraction (EBSD) in conodonts. Lethaia, Vol. 45, pp. 253–258. Conodonts are extensively used as ‘geochemical tools’ in palaeoenvironmental reconstructions in Palaeozoic and Triassic strata. The retention of a primary geochemical signal is substantially dependent on the ultrastructural integrity and crystallinity after diagenesis. Direct assessments of the preservation potential of the polycrystalline matrix are scarce with the exception of the application of transmission electron microscopy (TEM). The possible application of electron backscatter diffraction (EBSD) is introduced for well‐preserved Pennsylvanian platform conodont elements as a novel in situ technique based on successful previous results for biogenic carbonates. EBSD shows the presence of no diffraction in central regions of all studied elements of Idiognathodus and Adetognathus, with possible implications for geochemistry analysis. The absence of diffraction could be attributed to the existence of an amorphous phase, water (+OH) or a process of hypocalcification related to the development of ‘growth cavities’ in the lamellar crown tissue. Overall results indicate the suitability of using EBSD to extract crystallographic information from conodonts and potentially for other apatite (micro‐) fossils. □Conodont, diagenesis, EBSD, geochemistry, Pennsylvanian.     

Correlation Between Shine–Dalgarno Sequence Conservation and Codon Usage of Bacterial Genes

In this study, we analyzed the correlation between codon usage bias and Shine–Dalgarno (SD) sequence conservation, using complete genome sequences of nine prokaryotes. For codon usage bias, we adopted the codon adaptation index (CAI), which is based on the codon usage preference of genes encoding ribosomal proteins, elongation factors, heat shock proteins, outer membrane proteins, and RNA polymerase subunit proteins. To compute SD sequence conservation, we used SD motif sequences predicted by Tompa and systematically aligned them with 5′UTR sequences. We found that there exists a clear correlation between the CAI values and SD sequence conservation in the genomes of Escherichia coli, Bacillus subtilis, Haemophilus influenzae, Archaeoglobus fulgidus, Methanobacterium thermoautotrophicum, and Methanococcus jannaschii, and no relationship is found in M. genitalium, M. pneumoniae, and Synechocystis. That is, genes with higher CAI values tend to have more conserved SD sequences than do genes with lower CAI values in these organisms. Some organisms, such as M. thermoautotrophicum, do not clearly show the correlation. The biological significance of these results is discussed in the context of the translation initiation process and translation efficiency. Received: 22 June 2000 / Accepted: 18 October 2000     

浙江黄芝山剖面二叠-三叠系界线附近的牙形类序列及其地层对比

首次系统研究浙江黄芝山剖面二叠-三叠系界线附近的牙形类化石,共鉴定2属16种,并据此识别出5个牙形类化石带,从下至上依次为:Clarkina changxingensis changxingensis带,C.parasubcarinata带,C.chan-gxingensis yini-C.meishanensis zhangi带,C.meishanensis meishanensis带,Hindeodus parvus带。Hindeodus changxingensis,H.eurypyge,H.parvus分别在第15层、第18层底部、第18层中部依次出现,为确定该剖面二叠-三叠系界线在18-3层之底提供了依据。作者还讨论黄芝山剖面二叠-三叠系界线附近的牙形类序列与浙江煤山、伊朗剖面二叠-三叠系界线层牙形类序列的对比。     

The structure of some Middle Cambrian conodonts, and the early evolution of conodont structure and function

Elements presumed to belong to conodontophorids from the lower Middle Cambrian of Scania, Sweden, are investigated with emphasis on histology. The new genus Amphigeisina is described, with A. danica (V. Poulsen) as type species. The material also includes Hertzina? bisulcata Miiller and Proconodontus? sp. These forms, here termed proto-conodonts, appear to have grown by basal-internal accretion, unlike elements of the essentially post-Cambrian Conodontiformes, but partly similar to the Late Cambrian para-conodonts. A proposed model of the early evolution of conodonts involves the following points: (1) Conodonts were primarily external elements, secreted by an ectodermal epithelium. (2) An evolutionary trend during the Cambrian involved gradual retraction of the elements into pockets of the epithelium. (3) Elements of the Conodontiformes, herein termed euconodonts, were completely engulfed in epithelial pockets, which enabled them to grow holoperipherally and thus to assume more complex shapes that could be retained during growth. (4) The secreting epithelium adhered only to the basal body of the euconodont, the conodont proper being extruded from the pockets when the apparatus was in function.     

江苏下扬子区寒武系-三叠系牙形刺色变指标(CAI)图及其意义

本文对多年来积累的江苏下扬子区寒武系－三叠系牙形刺CAI材料进行了系统收集和整理,并采用Anita Harris建立的统一标准,编制了本区寒武系－三叠系的5幅牙形刺CAI图;进而结合牙形刺CAI与镜质组反射率（Ro)的对应关系,对上述各系的有机质成熟度进行了评价,对本区海相中、古生界油气勘探具有十分重要的意义。     

Ichnology and sedimentology of the early Permian deep-water deposits from the Lercara-Roccapalumba area (Western Sicily,Italy)

Summary Diverse and abundant trace fossils of the deep-waterNereites ichnofacies have been found in well-dated Early Permian deep-water turbidites (Lercara Formation) of western Sicily (Italy). Conodonts indicate a latest Artinskian to Cathedralian (late Early Permian) age. Microfossils (pelagic conodonts, albaillellid Radiolaria, paleopsychrospheric ostracods, foraminiferal associations dominated byBathysiphon), trace fossils (deep-bathyal to abyssalNereites ichnofacies) and sedimentologic data collectively indicate a deep-water environment for the Early Permian turbidites of the Lercara Formation. The dominance ofAgrichnium and of thePaleodictyon subichnogeneraSquamodictyon andMegadictyon suggests that this icnofauna is closely related in ichnotaxonomic composition to other late Paleozoic deep-water ichnofaunas. The occurrence ofAcanthorhaphe. Dendrotichnium andHelicoraphe, to date only reported from Cretaceous or Tertiary flysch deposits, suggests that the entire ichnofauna corresponds well to previously documented Silurian-Tertiary flysch ichnofaunas. Eight new ichnospecies and a new ichnosubgenus,Megadictyon, are described.     

Finite element,occlusal, microwear and microstructural analyses indicate that conodont microstructure is adapted to dental function

Conodonts constitute the earliest evidence of skeletal biomineralization in the vertebrate evolutionary lineage, manifest as a feeding apparatus of tooth‐like elements comprised of enamel‐ and dentine‐like tissues that evolved in parallel with these canonical tissues in other total‐group gnathostomes. As such, this remarkable example of evolutionary parallelism affords a natural experiment in which to explore the constraints on vertebrate skeletal evolution. Using finite element analysis, informed by occlusal and microwear analyses, we tested the hypothesis that coincidence of complex dental function and microstructural differentiation in the enamel‐like tissues of conodonts and other vertebrates is a consequence of functional adaptation. Our results show topological co‐variation in the patterns of stress distribution and crystallite orientation. In regions of high stress, such as the apex of the basal cavity and inner parts of the platform, the crown tissue comprises interwoven prisms, discontinuities between which would have acted to decussate cracks, preventing propagation. These results inform a general occlusal model for platform conodont elements and demonstrate that the complex microstructure of conodont crown tissue is an adaptation to the dental functions that the elements performed.     

New Middle Devonian conodont data from the Dong Van area,NE Vietnam (South China Terrane)

Middle Devonian conodonts from the Si Phai section in NE Vietnam are described. The section ranges from the Middle Devonian ensensis to timorensis conodont zones to the Late Devonian rhomboidea conodont Zone. A rich overall assemblage is described, including 27 taxa of species or subspecies rank and 11 taxa described in an open nomenclature. Among the dominant Polygnathus forms, four new taxa are described: Polygnathus linguiformis saharicus subsp. nov., Polygnathus linguiformis vietnamicus subsp. nov., Polygnathus rhenanus siphai subsp. nov., and Polygnathus xylus bacbo subsp. nov. Conodont assemblages are attributed to polygnathid, polygnathid-klapperinid, and klapperinid conodont biofacies representing hemipelagic to pelagic environments. The klapperinid biofacies, unreported in the previous literature, are here attributed to offshore areas of the external shelf. The taxonomic compositions of the studied conodont assemblages, as well as their CAI characteristics (CAI 4–5), suggest a palaeogeographic affinity of the studied strata to the Chinese Devonian Guangxi Basin, and the South China Terrane in general. Furthermore, the conodont biofacies and the palaeogeographic distribution of the fauna are discussed.