DEVONIAN CONODONT APPARATUSES AND THEIR VICARIOUS SKELETAL ELEMENTS

Recognition of four types of skeletal apparatuses of Devonian compound and platform conodonts, which conform in their basic plan to those already established in the Ordovician, Silurian, and Carboniferous, emphasizes the persistence of but a few structural types in the Paleozoic. The vicarious nature primarily of compound but also of platform elements in different multielement associations demonstrates mosaic evolution in Devonian apparatuses and has far-reaching taxonomic implications. Homologies of skeletal elements, for which a system of symbols is introduced, are evident in different apparatus types and show that the relationships of elements have been obscured by the older form taxonomy. Multielement analysis revises the taxonomic scope of the common Devonian genera, Icriodus, Polygnathus , and Ozarkodina. As a consequence of apparatus reconstruction, four new genera are proposed: Parapolygnathus, Cryptotaxis, Delotaxis , and Pedavis.     

ELEMENT ARRANGEMENT IN CONODONT APPARATUSES OF HINDEODELLA TYPE AND IN SIMILAR FORMS

The criteria used for logical and numerical reconstruction of conodont apparatuses are: (1) Similar stratigraphical and geographical occurrence. (2) Similar frequency variation. (3) Similar appearance of details in the conodont-elements. (4) Similar composition of the conodont apparatuses.
The position of conodont-elements in some types of apparatuses, as reconstructed by Lange (1968), is supported by further evidence, and it is also possible to reconstruct the precise position of certain elements. It is probable that a great number of apparatuses with compound conodont elements consisted of homologous elements and that a reconstruction of the positions of the elements from cluster material can be generalized to include all these apparatuses. From the periphery inwards, a complete apparatus of this type consisted of ne, hi, hi, pl, and tr elements and, posterior to these, of pairs of oz and sp elements.     

Parafurnishius,an Induan (Lower Triassic) conodont new genus from northeastern Sichuan Province,southwest China and its evolutionary implications

A fundamental aspect of taxonomy at the generic level, critical to understand Early Triassic conodont evolution, is the composition of the multielement apparatus. In this paper, we document a platform-bearing new conodont genus, Parafurnishius n. gen., as well as its multielement apparatus from the Griesbachian Feixianguan Formation (Lower Triassic) in Xuanhan County, northeastern Sichuan Province, southwest China. The new conodont genus is characterized by numerous robust and irregularly distributed conical denticles with variable platform morphology that has a possible affinity with the P₁ elements of Furnishius. These genera have apparatuses similar to those of Ellisonia and are classified with the family Ellisoniidae. The strong intraspecific variation of P₁ elements and the growth series within the entire sample population suggest that Parafurnishius may have evolved from the Griesbachian Isarcicella by developing random denticle positioning away from the platform centre, and then possibly evolved into younger Triassic Furnishius by developing a stable blade configuration. This preferred interpretation implies an ellisonid apparatus for Isarcicella. Alternatively, Parafurnishius may have evolved from Ellisonia and developed a homeomorphic P₁ element with Isarcicella. This new taxon has strong intraspecific variation of denticle growth orientation during the Early Triassic.     

Conodont affinity of the enigmatic Carboniferous chordate Conopiscius

Conopiscius shares V-shaped myomeres with the co-occurring conodont Clydagnathus but instead of a complex oral apparatus it has only a single pair of conical elements, and structures resembling scales are associated with its myomeres. Moreover, the coarsely crystalline crown tissue typical for conodonts has not been identified in the Conopiscius elements, which show only a finely lamellar skeletal tissue. The gap between conodonts and Conopiscius may be filled by isolated elements of similar morphology and structure occurring in the Late Devonian. They reveal a very thin external layer developed mostly at the tooth tip and resembling conodont crown tissue. The pulp cavity is partially filled with layered or spherulitic phosphatic tissue of the kind known also in conodonts (basal filling tissue) and early vertebrates (lamellin). Conodont elements of similar morphology and representing uni-membrate oral apparatuses have not been previously reported from the Devonian or Carboniferous but occur near the Cambrian–Ordovician transition ( Proconodontus ) and in the Late Permian ( Caenodontus ). It is proposed that Conopiscius represents a mostly cryptic conodont lineage extending from the Early Ordovician to the Permian, instead of being directly related to the agnathans.     

A 17‐element conodont apparatus from the Soom Shale Lagerstätte (Upper Ordovician), South Africa

Abstract:  Natural assemblages of a new conodont taxon, Notiodella keblon, from the Upper Ordovician Soom Shale Lagerstätte of South Africa contain 17 elements. This is the first time that a 17‐element apparatus plan has been unequivocally demonstrated in conodonts. The apparatus comprises paired P₁, P₂, P₃, M, S₁, S₂, S₃ and S₄ elements and an unpaired, axial S₀ element and provides a new template for use in the reconstruction of apparatuses from the collections of dispersed elements, particularly for those with icrion‐bearing P₁ elements and perhaps for other balognathids.     

A new Middle Ordovician conodont from central Kazakhstan, nothern Kyrgyzstan, and Altai

Naimanodus degtyarevi gen. et sp. nov., a new conodont genus and species from the Lower Darriwilian (Middle Ordovician) limestones of the Naiman Formation in central Kazakhstan is described. Three types of coniform nongeniculate elements are identified in the apparatus of this species. The new species was widespread in the Asian Paleobasin and recorded in three localities in Kazakhstan, eastern Kyrgyzstan, and Altai.     

A NEW EARLY ORDOVICIAN CONODONT GENUS FROM THE SOUTHERN MONTAGNE NOIRE, FRANCE

Abstract:  Two species of a new Tremadocian (Early Ordovician) conodont genus from the Saint Chinian Formation of the southern Montagne Noire, France, are erected: Hammannodus sarae gen. et sp. nov. and Hammannodus juliae gen. et sp. nov. They were found within a single storm-induced limestone nodule interbedded with offshore shales belonging to the regional Shumardia ( C .) pusilla (trilobite) Biozone, and to the Paltodus deltifer deltifer (conodont) Subzone. This conodont record is associated with the episodic development of carbonate productivity in temperate waters of the Montagne Noire platform, a process absent in neighbouring platforms of north-west Gondwana. The apparatus is composed of five coniform pyramidal elements occupying P and S positions and one bicostate element in the M position, having three or two sharp costae, respectively, with a subtriangular basal outline.     

Provinces, communities, and the proposed nektobenthic habit of Ordovician conodontophorids

Ordovician conodontophorid distribution, paleogeography, paleoclimate, and global tectonics strongly support the existence of one biogeographic conodontophorid province, largely restricted to equatorial regions and characterized by raised salinity and temperature. This Midcontinent Province is recognized in North America, Australia, and Siberia. The same data indicate that in other areas the North Atlantic Province represents a normal-marine, virtually cosmopolitan fauna. In both provinces, communities are recognized that show a lateral segregation, strongly indicating that the majority of Ordovician conodontophorids were benthic or nektobenthic. The only Ordovician pelagic forms we recognize are the simple cone genera that either lack, or have a simple, symmetry transition series. It appears possible to correlate certain evolutionary changes in the conodont apparatus with changes in preferred habitat.     

Morphofunctional analysis of Late Paleozoic conodont elements and apparatuses

Five functional types are recognized among Late Paleozoic conodont elements; they are distinguished by morphological and histological characteristics: grasping-holding, filtering, cutting, crushing, and grinding. Combinations of functional types of elements form functional types of conodont apparatuses, the main of which are filtering, grasping-cutting, grasping-pressing, grasping-cutting-grinding, and grasping-cutting-crushing apparatuses.     

AnOistodus venustus-like conodont species from the Middle Ordovician of Baltoscandia

Ordovician conodont specimens resemblingOistodus venustus Stauffer, 1935 have been reported from many areas. There is increasing evidence, however, that several lineages with homeomorphic conodont elements have erroneously been referred to one and the same species. I have investigated Baltoscandian conodont elements of this kind in order to find out about their origins and phylogenetic relationships with morphologically comparable elements from other areas. A natural grouping of finds from the Middle and lower Upper Ordovician of Baltoscandia is here described as belonging to a new species of a new genus,Venoistodus balticus n. gen., n. sp. The new species probably evolved paedomorphically fromDrepanoistodus forceps (Lindström, 1955) in the Early Ordovician.     

EARLY ORDOVICIAN CONODONTS FROM TARUTAO ISLAND,SOUTHERN PENINSULAR THAILAND

Abstract: Early Ordovician conodont faunas of the Thung Song Formation on Tarutao Island, southern peninsular Thailand, consist of 14 known species belonging to 17 genera, and eight undescribed species. Utahconus tarutaoensis and Filodontus tenuis are new species. Three conodont zones: the Rossodus manitouensis Zone, the Utahconus tarutaoensis Zone and the Filodontus tenuis Zone, in ascending order, are defined in the study sections. These are coeval with the interval from the Rossodus manitouensis Zone to the Acodus deltatus‐Oneotodus costatus Zone of the standard zonation in the North American Midcontinent. Based on the conodonts studied here and fossils previously reported from Tarutao Island, the Thung Song Formation is early Tremadocian to middle Arenig (Ibexian) in age. This formation is lithostratigraphically subdivided into the S1 to S5 members, and our study sections consist of the S1 to S3 members. These strata accumulated on a shelf in the Early Ordovician. The depositional environments of the limestones making up the S1 and S3 members were in deeper‐shelf conditions. Limestone and shale of the S2 member formed in a shallow‐water, high‐energy environment.     

Middle Triassic conodont apparatus architecture revealed by synchrotron X-ray microtomography

The composition of conodont apparatuses is crucial for understanding the feeding mechanisms of these early vertebrates. However, the multielement apparatus reconstructions of most species remain equivocal because they have been inferred from loose element collections, guided by knowledge from rare articulated ‘bedding plane assemblages’ and fused clusters, often from distantly related taxa. Even these natural assemblages can be difficult to interpret because the component elements can be closely juxtaposed or embedded in matrix, making it hard to discern the morphology of each element and their relative positions within the architecture of the feeding apparatus. Here we report five exceptionally preserved conodont clusters from the Middle Triassic Luoping Biota, Yunnan Province, Southwest China. These materials were scanned using synchrotron radiation X-ray tomographic microscopy (SRXTM), revealing the morphology and positional homology of the component elements in the fused clusters. We confirm that the apparatus of Nicoraella was composed of eight types of elements, comprising a total of 15 elements. SRXTM reveals the positional homologies of the component elements, viz. a single alate element is located in the S₀ position, flanked successively abaxially by pairs of breviform digyrate S₁ and S₂ elements, bipennate S₃ and S₄ elements, and a pair of inwardly curved breviform digyrate M elements. Carminate elements occupy the P₁ and P₂ positions. The apparatus of Nicoraella is among the most completely characterised of all conodonts and serves as a template for the reconstruction of gondollellids.     

A spathognathodont lineage of Mississippian conodonts

Evolutionary lineages of conodont elements and of some apparatuses are gradually being documented to increase the value of these fossils in stratigraphic interpretation and to further our understanding of conodont taxonomy. The well-documented faunas from Mississippian strata of the Illinois Basin and type Mississippian beds provide a great deal of information for the recognition of phylogenies. The disjunct element genus Spathognathodus is part of a basic and long-ranging type of conodont apparatus from which a number of other forms have been derived, and some of its Mississippian disjunct element species can now be placed in multielement species. A lineage involving spathognathodont Pa elements recognized in the Mississippian, then, is the basis of the present study. The phylogeny recognized started with Synprioniodina? crassidentata, and includes S.? regularis, S.? pulchra, S.? coalescens, S.? spicula, Hinduodus cristula, and H. minutus in the Mississippian. Aputiignathus scitulus and A. penrscitulus apparently represent a separate line, which also might include disjunct element Spathognathodus rexroadi.     

The new chronostratigraphic classification of the Ordovician System and its relations to major regional series and stages and to δ13C chemostratigraphy

The extensive work carried out during more than a decade by the International Subcommission on Ordovician Stratigraphy has resulted in a new global classification of the Ordovician System into three series and seven stages. Formal Global Boundary Stratotype Section and Points (GSSPs) for all stages have been selected and these and the new stage names have been ratified by the International Commission on Stratigraphy. Based on a variety of biostratigraphic data, these new units are correlated with chronostratigraphic series and stages in the standard regional classifications used in the UK, North America, Baltoscandia, Australia, China, Siberia and the Mediterranean‐North Gondwana region. Furthermore, based mainly on graptolite and conodont zones, the Ordovician is subdivided into 20 stage slices (SS) that have potential for precise correlations in both carbonate and shale facies. The new chronostratigraphic scheme is also tied to a new composite δ¹³C curve through the entire Ordovician.     

A Light and Scanning Electron Microscopic Study of the Closing Apparatus in Tintinnid Ciliates (Ciliophora,Spirotricha, Tintinnina): A Forgotten Synapomorphy

ABSTRACT. A membranous closing apparatus shuts the lorica opening in disturbed tintinnids of six genera belonging to four families. The homology of the apparatuses is investigated, using data from the literature and Mediterranean tintinnids studied in vivo and by scanning electron microscopy. Morphological and functional similarities indicate that the foldable closing apparatus is not only a synapomorphy of the genera Codonella (Codonellidae) and Dictyocysta (Dictyocystidae), as suggested 80 years ago, but also of Codonaria (Codonellidae) and Codonellopsis (Codonellopsidae). In Codonaria, Codonella, and Dictyocysta, the apparatuses merge posteriorly into membranous lorica sacs, which probably represent homologous structures. The diagnoses of these genera are improved according to the new findings. The close relationship of Codonella, Codonellopsis, and Dictyocysta is also inferred from small subunit rRNA phylogenies and the ultrastructure of the capsules. It contradicts the current lorica‐based classification of the tintinnids. The assumption that the diaphragm‐like apparatus in the genera Salpingacantha and Salpingella is not homologous to the foldable ones in the genera mentioned above is supported by molecular and cytological features.     

Zur Biologie, Taxonomie und Chronologie der Conodonten

The actual state of knowledge concerning the conodont animal is critically reviewed. All soft tissue documented by the specimens of the conodont animal were interpreted by the British collectors as Craniota (Vertebrata) features. Opinions on the precise affinities, however, remain controverse: The alleged chorda could have been a gut, bilateral symmetry, myomery, apatitic biomineralisation occur also in Invertebrata, caudal fins do in Chaetognatha. Even the lobes in the capital area interpreted as sensory organs could have served as aiding organs for food intake. Therefore, the originally sensational findings are reduced to the core issue that the conodont animal had an eel-like body and at the blunt end of it, behind peculiar lobes, the mineralized conodont elements were arranged. As a consequence, a relationship to the Protochordata cannot be excluded. A systematic position of the conodont animal as possible Vertebrata is likely only if organized below the Ostracodermata which is particularly so because conodonts occur so early in the Cambrian. In the second part of the article conodonts are discussed as guide fossils. The comprehensive, empirically grown taxonomy of the single elements is the prerequisite for recognizing outstanding, high resolution stratigraphy. In comparison, a multielement taxonomy remains as long problematical, as it is based on assumed, mostly statistically reconstructed rather than on natural multielement apparatuses. Taxonomical and nomenclatorial operations built on assumed apparatus affinity can bear heavily on nomenclature and stratigraphy of platform elements. It can be shown at the example of the Devonian that the platform elements have provided particularly in the pelagic facies a Standard Conodont Zonation that covers time almost without any gaps. The basis for this is that platform single element species and their phylogenetic development are extra-well known. In a special paragraph, history and applicability of the Standard Zonation is discussed in details. Different conodont biofacies are developed between the pelagic realm where the Standard Zonation originated, and the coastal areas so that conodonts can indicate temporal and bathymetric relationships of the mother rocks. In addition alternative conodont zonations were introduced mainly for the shallow water. Because of their conspicuous stratigraphic significance in the Devonian, conodonts were employed by International Committees to redefine Series and Stage Boundaries. For the boundary selection species were utilized from phylognetic lineages as only from tight evolution their point of origination could be well identified and recognized in worldwide correlation. The GSSPs (Global Stratotype Section and Point) within the Devonian are discussed in detail. In recent times, attempts are made at calibrating the duration of conodont zones on the basis of their well known rate of evolution and high temporal resolution. These analyses have to be continued in order to demonstrate the absolute time relationship of condont zones in more and continuous detail. Currently the phyletically controlled Standard Conodont Zonation is the authoritative stratigraphic method in the Devonian. It has provided together with the GSSPs precise and valuable fix points for a Global Time Scale (GTS). In combination with additional stratigraphie methods the Standard Conodont Zonation may provide further refinements in the future.     

Cutting the first ‘teeth’: a new approach to functional analysis of conodont elements

The morphological disparity of conodont elements rivals the dentition of all other vertebrates, yet relatively little is known about their functional diversity. Nevertheless, conodonts are an invaluable resource for testing the generality of functional principles derived from vertebrate teeth, and for exploring convergence in a range of food-processing structures. In a few derived conodont taxa, occlusal patterns have been used to derive functional models. However, conodont elements commonly and primitively exhibit comparatively simple coniform morphologies, functional analysis of which has not progressed much beyond speculation based on analogy. We have generated high-resolution tomographic data for each morphotype of the coniform conodont Panderodus acostatus. Using virtual cross sections, it has been possible to characterize changes in physical properties associated with individual element morphology. Subtle changes in cross-sectional profile have profound implications for the functional performance of individual elements and the apparatus as a whole. This study has implications beyond the ecology of a single conodont taxon. It provides a basis for reinterpreting coniform conodont taxonomy (which is based heavily on cross-sectional profiles), in terms of functional performance and ecology, shedding new light on the conodont fossil record. This technique can also be applied to more derived conodont morphologies, as well as analogous dentitions in other vertebrates and invertebrates.     

Patterns of bilateral asymmetry and allometry in Late Devonian Polygnathus conodonts

Conodont animals were early jawless vertebrates equipped with a feeding apparatus composed of several tooth‐like elements. The P1 elements, at the rear of the apparatus, were characterized by a robust shape and rapid morphological evolution. Occlusion occurred between paired right and left P1 elements, occasioning some bilateral asymmetry, which, together with allometric growth, may partially obliterate the temporal differences. The present study aims to disentangle these different components of morphological variation in Late Devonian Polygnathus P1 conodont elements. An extensive 2D geometric morphometric analysis of the platform shape was performed through the Famennian record of two outcrops. This analysis was completed by a 3D study on a subset of conodont elements. The 2D and 3D morphometric quantifications provided highly congruent results, showing that the 2D shape constitutes a good approximation of the element geometry. The 3D analysis delivered further insights into the relationship between the geometry of the elements and the constraints related to occlusion. The 2D analysis allowed a quantitative assessment of the variation among species and through time. Allometry and bilateral asymmetry were differently expressed depending on the species considered, suggesting that constraints imposed on pairing by the morphology of the elements varied even among related species. The within‐species variation was so important that it largely obliterated temporal trends; a relationship of Polygnathus shape and conodont biofacies variations through the Famennian nevertheless suggested an evolution driven by ecological interactions between conodont genera.     

Apparatus Composition, Growth, And Survivorship Of The Lower Ordovician Conodont paracordylodus Gracilis Lindström, 1955

Analysis of numerous conodont element clusters from the Lower Ordovician cherts of the Burubaital Formation in central Kazakhstan reveals that the apparatus of Paracordylodus gracilis Lindstro¨m, 1955 consisted of 15 elements: two M elements, nine S elements (including 1 S₀), and four P elements (2 P₁, 2 P₂). The clusters probably originated as faecal pellets, but the best preserved indicate that the architecture of the apparatus of P. gracilis was comparable to that of ozarkodinid conodonts, providing strong support for the hypothesis that the 15-element 2M-9S-4P apparatus plan was plesiomorphic for conodonts with morphologically complex elements. All the elements within the P. gracilis clusters appear to be at a similar stage of ontogeny, and there is no evidence for late addition or replacement of elements. Analysis of element growth suggests that the relative dimensions of some elements changed during ontogeny, but the available data support the hypothesis that the growth of the apparatus as a whole was isometric. The size distribution of P. gracilis in the Burubaital Formation suggests that individuals in a particular size range were preferentially selected for consumption by predators. The identity of these predators is unknown, but they may have included other P. gracilis .