The structure and evolution of Ordovician conodont apparatuses

Multielement taxonomy was instituted for Ordovician conodonts over a decade ago, and probably a majority of the multielement genera have been defined or are well understood. The present systems of notation for elements within apparatuses are inadequate and cumbersome. A new notation scheme is proposed which applies a single-letter code to the position in the apparatus occupied by certain element morphotypes. The taxonomic status of all known Ordovician conodont genera is reviewed (appendix) using the new notation, and a new scheme to classify conodont apparatuses is presented. Five main apparatus types (I-V) and seventeen subtypes (IA-IC, etc.) are defined. Within these groups, all known Ordovician conodont genera can be accommodated, and probably few new groups are required to include all other conodont genera. The apparatus types and subtypes are defined on the basis of symmetry, curvature, and number of the element types, with a clear distinction being made between the first and second transition series. Certain homologous relationships, both between and within many apparatus types, are noted. The evolution of the five major types, and the subtypes, is traced through the Ordovician. The pattern of evolution suggests that the types and subtypes recognized are probably natural biologic groupings, largely reflecting phylogenetic change.     

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.     

Early Triassic conodont clusters from South China: revision of the architecture of the 15 element apparatuses of the superfamily Gondolelloidea

Abstract: Several fused clusters of conodont elements of the genera Neospathodus and Novispathodus were recovered from limestone beds at the Dienerian–Smithian and Smithian–Spathian boundaries, respectively, from several localities in Guangxi province, South China. Conodont clusters are otherwise extremely rare in the Triassic, and these are first described for the Early Triassic. The exceptional specimens partially preserve the relative three‐dimensional position and orientation of ramiform elements and are therefore extremely important for testing hypotheses on the architecture of apparatuses. These specimens partly confirm the previous reconstruction of the Novispathodus apparatus by Orchard. Within apparatuses of members of superfamily Gondolelloidea, elements previously identified as occupying the S₁ and S₂ positions instead occupy the S₂ and S₁ positions. Similarly, within apparatuses of members of the subfamily Novispathodinae, elements previously referred to S₃ and S₄ positions are reinterpreted to have occupied S₄ and S₃ positions, respectively.     

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.     

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.     

Gondolelloid multielement conodont apparatus (Scythogondolella) from the Lower Triassic of Jiangsu,East China,revealed by high-resolution X-ray microtomography

The application of high-resolution X-ray microtomography on conodont natural assemblages has provided new information on the structure of the elements and enabled the three-dimensional reconstruction of apparatuses. We herein report four conodont natural assemblages from uppermost Lower Qinglong Formation, Longtan, Nanjing, East China. Using X-ray microtomography, we reconstructed the apparatus of the end-Smithian (Olenekian, Early Trassic) Scythogondolella milleri. Our result confirms that Scythogondolella has an octomembrate apparatus composed of 15 elements (a single S₀ element, two pairs of S_1–4, M and P_1–2 elements) like other gondolelloid apparatuses that have been tested by their corresponding natural assemblages, including Neogondolella, Novispathodus and Nicoraella. Element morphology of Scythogondolella closely resembles that of Neogondolella apparatus from the Illyrian (Anisian, Middle Triassic) of Monte San Giorgio: an alate (hibbardellan) S₀ element with two long lateral processes that meet at a denticle anterior of the cusp, a bipennate (hindeodellan) S₃ element with a bifurcated anterior process branching from the third denticle anterior of the cusp and an enantiognathiform S₂ element with two dissimilar processes of nearly equal in length. It differs from the latter in the length of the inner-lateral process of M element and the posterior process of S₀ and S_3–4 elements without considering the obvious morphological differences in P elements between them. The element positional homologues of Scythogondolella conforms to those of the standard 15-element plan shared primitively among ozarkodinin, prioniodinin and prioniodontid conodonts, and again confirms that the breviform digyrate elements of cypridodellan and enantiognathiform morphotypes occupy the S₁ and S₂ positions, respectively, within the superfamily Gondolelloidea.     

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.     

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 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.     

Eine Conodontengruppe von Prooneotodus tenuis (MÜller, 1959) in natürlichem Zusammenhang aus dem Oberen Kambrium von schweden

A natural conodont assemblage,Prooneotodus tenuis (MÜller, 1959) was discovered in shales in Zone I of the Upper Cambrian of Hunneberg, Väastergötland, Sweden. It is composed of 12 similar single cone elements, which form 6 pairs of different size. The largest is more than twice the size of the smallest. This variation in size together with the occurrence of between 8-12 elements in conspecific assemblages recorded by Miller & Rushton (1973) makes it likely that during growth the animal added new elements to the apparatus. Should this be applicable to all conodonts, this observation would be of some relevance in the statistical reconstruction of conodont apparatusses.     

Cladistic tests of monophyly and relationships of biostratigraphically significant conodonts using multielement skeletal data –Lochriea homopunctatus and the genus Lochriea

Abstract: Since the 1960s, huge progress has been made in reconstructing the multielement skeletons of conodont species and developing a biologically defensible taxonomy. Nevertheless, a widespread prejudice remains that certain parts of the conodont skeleton, particularly the P₁ elements, are more informative than others with regard to taxonomy and evolutionary relationships. Here, we test these views. A new partial multielement reconstruction of the skeleton of the biostratigraphically significant conodont originally described as Gnathodus commutatus homopunctatus allows us to conduct a cladistic test of the alternative hypotheses of phylogenetic placement of this species. Our analysis also provides the first test of the hypothesis that Lochriea– species of which are markers for global correlation – is monophyletic and tests hypotheses concerning the origins of the genus. Our results demonstrate that homopunctatus is a species of Lochriea and that the genus is monophyletic. The widely held view that Lochriea arose from a species of Bispathodus is not supported. Our results show that it is difficult to predict a priori which parts of the conodont skeleton carry phylogenetic signal, and provide strong support for the hypothesis that similarity in the morphology of conodont P₁ elements alone is not a reliable guide to relationships and taxonomic groupings of conodont species. This is because P₁ elements with similar morphologies are convergently acquired in multiple conodont clades, because reliance on the characters of only one of the six or seven morphologically distinct elements of the conodont skeleton ignores phylogenetically significant data and because P₁ elements can lack characters that might seem to be diagnostic of a genus. Conodonts are no different to other organisms: ignoring data that have the potential to be phylogenetically informative is unlikely to produce the most reliable hypotheses of evolutionary relationships. We suggest that other biostratigraphically significant hypotheses of relationship between conodont taxa that are based on P₁ elements alone should be subject to cladistic testing.     

Conodont assemblages: A new functional model

A functional model for the conodont apparatus is proposed based on Carboniferous bedding plane assemblages. It is proposed that the conodont assemblage functioned as a filter feeding and respiratory organ in organisms of unknown affinities, and that the organisation of elements in the assemblage genus Scottognathus can be used as a basic plan for interpreting the arrangement of conodont elements in other assemblages. In our model the Platform Blades opened and closed the anterior end of the system, the sieve consisted of Arched Blades, Pick-shaped Blades (assemblage genera Lewistownella and Lochriea) and the main cusps of the Elongate Blades; the denticulated bars of Elongate Blades and Pick-shaped Blades (form genera Scottognathus and Westphalicus) acted as a support for ciliated tissue. A radial and bilateral arrangement for the Elongate Blades is presented and discussed. Using this model as a guide we have reconstructed the assemblage genus Duboisella.The success of conodont organisms is attributed to the genetic plasticity of elements of the filter respiratory system, which enabled the system to adapt to evolutionary changes in the marine plankton. Ideas for further research are presented. In this account we have chosen to refer to the morphological form of conodont elements (Rhodes, 1954) rather than to the form generic and specific names assigned to them in the literature, e.g., we use the term Elongate Blades, instead of Hindeodella.     

Conodont biostratigraphy of the Early Triassic in eastern Slovenia

The Early Triassic is a critical interval for the study of recovery from the terminal Permian mass extinction, as there are small-scale extinction events, which may have contributed to the delayed recovery. The systematic measuring and sampling of a 12-m-thick section at the Mokrice locality in eastern Slovenia has resulted in the recovery of a conodont fauna from the Olenekian beds. Four conodont zones have been recognized. These zones are in ascending order as follows: the Hadrodontina aequabilis Zone, Platyvillosus corniger Zone, Platyvillosus regularis Zone, and Triassospathodus hungaricus Zone. These conodont zones confirm the proposed conodont biozonation sequence in western Slovenia and have correlation value especially for the western marginal Tethys. Multielement conodont apparatuses of Triassospathodus hungaricus and Platyvillosus regularis have been reconstructed based on conodont elements that were recently obtained from the Slovenian sections. Although the S₂element was not found, the apparatus indicates that the conodont species “Spathognathodus” hungaricus should be assigned to the genus Triassospathodus.     

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.     

The hydrodynamics of conodont elements

Accurate palacobiological interpretation of the fossil record requires an assessment of biostratinomic processes. Samples which have undergone significant post-mortem sorting reflect only the hydrodynamic regime prevalent at the time of deposition and not the original faunal composition. Conodont bedding plane assemblagcs are rare and many conodont taxa are known only through reconstructions of apparatuses using empirical techniques. These methods require large collections where the elements have undergone no significant post-mortem transportation. In this paper measurements of settling velocities are used to predict relative hydrodynamic behaviour in a current. The settling velocity of a particle is of about the same magnitude as the current required to entrain that particle. Conodont elements rotate during fall, spinning about a central axis. Elements with equal Reynolds numbers may have different drag coefficients indicating that shape is an important factor in determining conodont element hydrodynamics. In a current of increasing velocity the predicted order of entrainment of elements of the same equivalent size is as follows: Polygnathus S. Ancyrodella Pa. Polygnathus Pa. Palmatolepis Pa. Polygnathus Pb. lcriodus Pa. Conodont elements grow by accretion of lamellae; to the first approximation. length is a satisfactory indication of size. Length can therefore be used to predict the settling velocities of elements, and which elements will sort together. Measurement of element lengths allows a rapid estimation of the degree of post-mortem sorting undergone by a sample and therefore its suitability for use in palaeobiological analysis. □ Conodont hydrodynamics, biostratinomy, drag coefficient, settling velocity. sorting.     

THE GOLGI APPARATUS IN CHICK CORNEAL EPITHELIUM: CHANGES IN INTRACELLULAR POSITION DURING DEVELOPMENT

The intracellular position of the Golgi apparatuses in the basal cell layer of the corneal epithelium in embryonic and hatched chicks has been studied in the light microscope by impregnating the Golgi apparatus with silver. During two distinct periods in development the Golgi apparatuses in the basal cells shift from an apical to basal position. Each of these periods correlates in time with the appearance of an acellular collagenous matrix beneath the epithelium. Examination of the basal epithelial cells in the electron microscope confirms the intracellular shifts in position of the Golgi apparatus. The results suggest that the Golgi apparatus shifts to the basal cell pole of the corneal epithelium in order to excrete connective tissue materials into the developing corneal stroma.     

A neontological interpretation of conodont elements based on agnathan cyclostome tooth structire function, and development

Krejsa, R. J., Bringas, P. Jr. & Slavkin, H. C. 1990 10 15: A neontological interpretation of conodont elements based on agnathan cyclostome tooth structure, function, and development. Lethaia , Vol. 23, pp. 359–378. Oslo. ISSN 0024–1164.
Speculation about a conodont-cyclostome connection has led us to search for and establish a biological basis for various characteristic structures in conodont elements. Measurements of juvenile hagfish palatal and lingual teeth overlap those of representative conodont elements, demonstrating a size correspondence of conodonts with teeth of living vertebrates. When hagfish tooth histology is compared with internal and surface topography (SEM) of hagfish, keratinous teeth and mineralized conodont elements, microspaces and tubules similar to those found in hagfish functional tooth coverings and replacement elements are also found within the white matter' of conodont elements. It is provisionally suggested that the primary organic matrix of conodont elements could be keratin and/or keratin-related molecules, and that individual conodont elements could represent shed tooth coverings. The basal bodies' found in certain conodont elements could be replacement elements. These interpretations are contrary to several paradigms of orthodox conodontology. ▭ Agnatha, conodonts, cyclostomes, hagfish, keratin, paleo-biology, shedding, teeth .     

Siphonodella leiosa (Conodonta), a new unornamented species from the Tournaisian (lower Carboniferous) of Puech de la Suque (Montagne Noire,France)

A new conodont species, Siphonodella leiosa, is described from the lower Carboniferous pelagic limestones of the Montagne Noire (France), deposited in North Gondwana on a outer platform environment. Specimens were obtained from one level dated to the Siphonodella jii conodont Zone. The major difference from other siphonodellid conodonts known in this area is that the elements of this new species have a practically entirely smooth and unornamented platform, apart from the development of one or two low rostral ridge-like nodes. Similar morphologies were generally observed in shallow marine deposits of the same time frame from China, Russia and East and Central European areas. The new discovery reinforces the idea that ornamentation of siphonodellids is not only related to bathymetry, but that temperature could play an important role in the diversification and radiation of unornamented species during the Siphonodella jii conodont Zone.     

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.     

Conodont element function

There are close similarities between conodont elements and teeth both in general shape and in that they possess pointed tips and have expanded bases with more porous tissue. A number of examples of conodont elements which parallel specific kinds of tooth shapes and organizations are added to earlier known similarities. Both in teeth and in conodont elements the surface structures include cutting edges, striations, and barbs. The change in strength of the conodont denticles caused by the evolution of white matter is also shown to agree with a tooth function. On the other hand, the elements grew throughout the life of the animal by lamellae added to the surface. The solution of this paradox is found in the elements alternating between a growth phase and a functional phase. During growth the oral surface of the elements was enveloped in folds of secreting soft tissue. Structures henceforth termed burrs were formed at the contacts between the folds. Parts of the burrs evolved into cutting edges. Three different bite types occurred among the conodont elements. Many (all?) conodonts were predators which used their elements to seize and to process the food mechanically. The shape of the conodont elements cannot be used for conclusions regarding the affinities of the conodonts. Similarly, an identification of a fossil as belonging to the conodonts must be supported by other evidence than just shape.