Katian (Upper Ordovician) conodonts from Wales

Middle and upper Katian conodonts were previously known in the British Isles from relatively small collections obtained from a few localities. The present study is mainly based on 17 samples containing more than 17 000 conodont elements from an approximately 14‐m‐thick succession of the Sholeshook Limestone Formation in a road cut near Whitland, South Wales, that yielded a diverse fauna of more than 40 taxa. It is dominated by representatives of Amorphognathus, Aphelognathus/Plectodina and Eocarniodus along with several coniform taxa. Representatives of Decoriconus, Istorinus and Sagittodontina are reported from the Ordovician of UK for the first time. The fauna is a typical representative of the British Province of the Atlantic Realm and includes a mixture of taxa of North American, Baltoscandic and Mediterranean affinities along with pandemic species. Based on the presence of many elements of Amorphognathus ordovicicus and some morphologically advanced specimens of Amorphognathus superbus, the Sholeshook Limestone Formation is referred to the lower A. ordovicicus Zone. Most of the unit is also coeval with Zone 2 of the Cautleyan Stage in the British regional stage classification, and stage slice Ka3 of the middle Katian Stage in the global stratigraphical classification, an age assignment consistent with data from trilobites, graptolites and chitinozoans. The unusually large collection of M elements of Amorphognathus provides insight into the complex morphological variation in this element of some Katian species of this genus. The Sholeshook conodont fauna is similar to those of the Crûg and Birdshill limestones, but differs in several respects from the slightly older ones from the Caradocian type area in the Welsh Borderland. Although having some species in common, the Sholeshook conodont fauna clearly differs from coeval Baltoscandic faunas and is even more different in composition compared with equivalent North American Midcontinent faunas.     

RECONSTRUCTION OF AN APPARATUS OF NEOSTRACHANOGNATHUS TAHOENSIS FROM ORITATE,JAPAN AND SPECIES OF NEOSTRACHANOGNATHUS FROM OMAN

Abstract: The apparatus of an Early Triassic conodont Neostrachanognathus tahoensis Koike, 1998 from Oritate, Kumamoto Prefecture, Japan, and a species of Neostrachanognathus from Oman were reconstructed. On the basis of five natural assemblages from the Oritate area, the three‐dimensional apparatus model of N. tahoensis is interpreted as bilaterally symmetrical and composed of 14 elements consisting of pairs of P₁, P₂, P₃, S₁, S₂, S₃, and S₄ elements. The P₁ and P₂ elements are coniform elements, the P₃ elements are digyrate forms, and the S elements are bipennate ramiforms. The S elements are arranged rostrally in the apparatus and the pairs of the P₁, P₂, and P₃ elements are subvertically arranged caudally and ventrally to the S array. One of the natural assemblages was formed by rostrocaudal collapse of the apparatus on the sea floor, whereas the other assemblages indicate that conodont animals came to rest nearly parallel with the substrate prior to burial. A collection of isolated elements from Jabal Safra, Oman, includes a second species of Neostrachanognathus with a comparable apparatus.     

Conodont chronostratigraphical resolution and Declinognathodus evolution close to the Mid‐Carboniferous Boundary in the Barcaliente Formation type section,NW Spain

The Barcaliente Formation in its type section provides significant conodont occurrences in the Mid‐Carboniferous Boundary interval (between the Mississippian and Pennsylvanian sub‐periods). A sequence of Declinognathodus morphotype appearances (considered here as species) is recognized from the uppermost Serpukhovian to the lower Bashkirian. These morphotypes belong to Declinognathodus noduliferus (Ellison & Graves) sensu lato, the primary marker for the worldwide correlation of the Mid‐Carboniferous Boundary. Among them, D. inaequalis (Higgins) appears to have been the first species to occur at the Mid‐Carboniferous Boundary bed in the Arrow Canyon GSSP. A morphocline between D. bernesgae (Sanz‐López et al.) and D. inaequalis is studied here to identify the bed correlated with the GSSP. Other diagnostic first occurrences of conodonts (Rachistognathus minutus (Higgins & Bouckaert), D. noduliferus, D. lateralis (Higgins & Bouckaert), Idiognathoides asiaticus Nigmadganov & Nemirovskaya and I. corrugatus (Harris & Hollingsworth)) are evaluated as chrono‐markers throughout their worldwide distribution, particularly for the Eurasian faunas. The available secondary markers provide a better resolution and support chrono‐correlation in sections other than that at the GSSP. The worldwide first and late occurrences of studied taxa are considered within the background of the Mid‐Carboniferous Extinction Event and the conodont overturn, where crises progenitor Declinognathodus gave rise to members of the Family Idiognathodontidae. Firstly, progenitor taxa are good biostratigraphic tools, whereas local extinction of genera considered Mississippian in age and new taxa arising later in the post‐crises radiation suggest an early Bashkirian palaeogeography and climatic conditions confining temporarily the biogeographic distribution of conodonts.     

Natural assemblages of Hindeodus conodonts from a Permian–Triassic boundary sequence,Japan

Hindeodus parvus and Hindeodus typicalis occur in a deep‐water chert and claystone section in the Mino Terrane, Japan, which has been identified as a Jurassic accretionary complex. Conodont fossils are preserved as natural assemblages of impression fossils on bedding planes in claystone. In this study, 13 assemblages of Hindeodus species were recognized, comprising at most 13 elements which generally maintain the original composition and structure of an apparatus. We discriminated pairs of carminiscaphate P₁, angulate P₂ and makellate M elements, as well as a single alate S₀ element and two digyrate and four bipennate elements constituting the S array. Although the digyrate and bipennate elements are preserved in the S₂ and S_3–4 positions, respectively, a pair of S₁ elements was not found due to incompleteness in the natural assemblages. The conodont biostratigraphy indicates that the lithological boundary between chert and claystone units in the study section corresponds exactly to the Permian–Triassic boundary.     

Testing hypotheses of element loss and instability in the apparatus composition of complex conodonts: articulated skeletons of Hindeodus

Knowledge of the conodont skeleton, in terms of the morphology of the elements and the positions they occupy, provides the foundation for understanding of homology, taxonomy and evolutionary relationships in conodonts. This knowledge also underpins analyses of conodont functional morphology and feeding. Direct evidence of skeletal anatomy and apparatus architecture comes from natural assemblages: fossils that preserve together the articulated remains of the conodont apparatus, either collapsed onto a bedding plane or as clusters of elements in which juxtaposed and overlapping elements have been fused together by diagenetic minerals. Here we describe six clusters of the biostratigraphically important conodont Hindeodus parvus from the Lower Triassic Shangsi section, Sichuan Province, South China. Five of these clusters represent the partial remains of articulated skeletons, providing direct evidence of the number and arrangement of elements in the apparatus. Combined with data from previously published natural assemblages this provides a test of the hypothesis that Triassic conodonts had a reduced dentition. Hindeodus parvus possessed a complete raptorial array of two M and nine S elements (unpaired S₀; symmetrically paired S₁, S₂, S₃, S₄); the paired P₁ locations were occupied by carminiscaphate elements, but the apparatus lacked P₂ elements. This is consistent with broader evidence for a particularly high degree of integration and constraint operating on the S–M array of morphologically complex conodonts, leading to conserved architecture of the array over a period of more than 250 million years. The loss of elements from the P domain implies a change in food processing ability and, given the predominance of data from P elements in conodont taxonomy and biostratigraphy, the hypothesis of element loss from the P domain has significant implications for the broader understanding of conodont diversity and evolutionary patterns.     

Conodont biostratigraphy of Tournaisian shallow-water carbonates in central Guangxi,South China

The conodont faunas of Tournaisian shallow-water carbonates from central Guangxi are described mainly for biostratigraphic purposes. A complete series of samples was collected from the Long’an and Du’an formations in the Long’an section. These formations are characterized by lime-mudstone, skeletal and peloidal wackestone, packstone and grainstone with typical shallow-water biota. Overall, these samples produced 809 identifiable Pl elements, belonging to 50 species in 11 genera, of which one species and one subspecies are new. The fauna enables the establishment of seven biozones (in ascending order): Polygnathus spicatus, Siphonodella homosimplex, S. sinensis, S. dasaibaensis, Polygnathus communis carina Acme, Gnathodus cuneiformis, and P. communis porcatus zones. Based on these new collections from central Guangxi and on data from the literature, a conflated Tournaisian conodont zonation is proposed for shallow-water successions in South China. Most of the conodont zones correlate well with their counterparts recognized in Western Europe, which may be of greater significance in stratigraphic correlation than previously thought.     

New insights into pollen morphology and its implications in the phylogeny of Sanguisorba L. (Rosaceae; Sanguisorbeae)

Pollen morphology of 13 species of the genus Sanguisorba (Rosaceae) was examined by light and scanning electron microscopy. The pollen morphology divided the genus into two main groups: (A) tricolporate with tapered colpus tips, and (B) either tricolporate with opened colpus tip or hexacolporate. The former group was further subdivided into those without vestibulum (A1: S.?alpina, S.?dodecandra, and S.?filiformis) and those with vestibulum (A2: S.?agrimonoides, S.?ancestroides, S.?annua, S.?cretica, S.?minor, and S.?verrucosa), whereas the latter group was subdivided into those with colpus narrow and similar to the A1 type (B1: S.?canadensis and S.?diandra), those with colpus and mesocolpus somewhat equally wide and forming hexacolporate aperture (B3: S.?albiflora, S.?armena, S.?media, S.?menziesii, S.?parviflora, S.?stipulata, and S.?tenuifolia), and those with colpus intermediate between the B1 and B3 types (B2: S.?hakusanensis, S.?microcephala, S.?obtusa, S.?officinalis, S.?polygama, and S.?sitchensis). It is suggested that the A1 type aperture would have evolved to the A2 type as a specialized form and the B types (B1 to B3) in a direction in which the hexacolporate aperture was derived. Implications of pollen morphology for infrageneric classification of Sanguisorba are discussed and the results are compared with molecular phylogenetic studies.     

A Tourist element in the 5′-flanking region of the catalase gene CatA reveals evolutionary relationships among Oryza species with various genome types

Tourist-OsaCatA, a transposable element, was found in the 5′-flanking region of the rice gene CatA. The characteristics of this element are similar to those of the other Tourist elements so far found in Oryza sativa. PCR and sequence analyses of 37 accessions of 18 species revealed that all the Oryza species examined, except for one accession, have either a full-length or a partial Tourist element at this locus. Unlike the Tourist elements previously reported, this Tourist element is found in all four Oryza species complexes in the Oryzeae tribe. All AA genome Oryza species, except O.?longistaminata, contain the full-length Tourist element. O. longistaminata and the species of the O. officinalis, O. meyeriana and O.?ridleyi complexes contain the partial element. A phylogenetic tree of Oryza species based on the nucleotide sequences of these Tourist elements was constructed. The O.?longistaminata accessions were placed near the neighboring cluster of the officinalis complex. We propose that the ancestor of O.?longistaminata and that of other species with the AA genome diverged, and the ancestor(s) of the O.?officinalis, O.?ridleyi and O.?meyeriana complexes then diverged from the ancestor of O.?longistaminata in the course of the evolution of the Oryza species. The Tourist elements associated with CatA and its orthologs thus provide useful tools for examining evolutionary relationships among Oryza species.     

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.     

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.     

SPORE TETRAD STRUCTURES OF POSSIBLE HEPATIC AFFINITY FROM THE AUSTRALIAN LOWER CARBONIFEROUS

Palynological studies of Early Carboniferous (Viséan) sediments of the Bonaparte Gulf Basin, northwestern Australia, reveal the presence of intact tetrahedral spore tetrads (here described as Sagenotetradites gen. nov.) and, more usually, of their disjunct spore portions which had previously been interpreted as dispersed whole-miospore species. Two species of the new tetrad genus are recognized: S. bonapartensis (Playford) comb, nov., as type species; and S. spiritensis (Playford) comb. nov. The species are distinguished, as originally, on the basis of sculptural attributes of their distal exoexines. When intact, both species share a common internal “binding” element (“Cadiospora abrupta” Playford, 1971) composed of the fused proximal exoexinal faces of all four spores of a given tetrad. Morphological comparisons with modern hepatic spores suggest an alliance of the microfossils with the order Sphaerocarpales. The occurrence of Sagenotetradites in exclusively marine sediments suggests that its parent plants grew in close proximity to the marine depositional basin. Moreover, the morphological attributes of the tetrads would appear to have facilitated dissemination by water.     

Is the serpentine aster, <Emphasis Type="Italic">Symphyotrichum depauperatum</Emphasis> (Fern.) Nesom,a valid species and actually endemic to eastern serpentine barrens?

Serpentine aster, Symphyotrichum depauperatum (Fern.) Nesom, is the ‘flagship’ species of the eastern serpentine barrens, inhabiting 20 of the 26 remaining occurrences of significant size of this globally rare community type and long recognized as its only known endemic species. Previous studies have called into question both the validity of the taxon and its status as a true endemic of the serpentine barrens. We used amplified fragment length polymorphism (AFLP) analysis to compare seven serpentine barrens populations, one alleged diabase glade population, and two populations each of the two species with which S. depauperatum is lumped by some authors. Our analysis supports the validity of S. depauperatum as a distinct species, which grows almost entirely on shallow soils overlying serpentinite bedrock in Pennsylvania and Maryland, but it confirms an earlier hypothesis that S. depauperatum also includes small, disjunct populations on diabase glades in North Carolina.     

Reproductive biology and isolation mechanisms in rupicolous species of the Acianthera prolifera complex (Orchidaceae) occurring in southeastern Brazil

We studied the floral biology and performed experimental intra- and interspecific pollinations in populations of a complex of four Acianthera (Orchidaceae) species occurring in Brazilian campo rupestre vegetation (A.?hamosa, A.?limae, A.?modestissima, and A.?prolifera). All four species flower synchronously, are partially intercompatible, and exhibited some degree of self-sterility. Floral morphology is similar in all the species, with their principal differences associated with size of the floral structures. The four species were visited only by Diptera species of the families Phoridae (Megaselia spp.) and Chloropidae, but visits are rare and fruit set is very low. Sympatric species were not pollinated by the same Diptera species. Acianthera hamosa and A.?modestissima have the smallest flowers, and no marked morphological differences between them were observed; they were both pollinated by very similar Megaselia species. Both prepollination barriers and postpollination events are important to maintaining the isolation of the species, functioning as overlapping filters that diminish the possibility of gene flow between them. However, putative hybrids between A.?prolifera and A.?limae have been found. Conversely, A.?hamosa and A.?modestissima, which are recognized only by vegetative characters that show high phenotypic plasticity, seem only to be isolated by geographical barriers, and they may actually constitute a single species or be sister species.     

A gnathodont lineage of Mississippian conodonts

Representatives of the disjunct element conodont genus Gnathodus described from lower Mississippian strata (Kinderhookian and Osagean Series) in the Midcontinent of North America illustrate a phylogenetic association composed of ten species. The lineage of G. delicatus — G. punctatus - G. semiglaber , of late Kinderhookian to early Osagean age, has been documented previously. Three distinct forms, G. cf. bilineatus, G. anfetexanus , and G. bulbosus , descended from G. semiglaber in early Osagean time. The latter two continued as parallel lineages; G. bulbosus - G. pseudosemiglaber and G. antetexanus - G. taxanus . In addition, two dead-end lineages, represented by G. typicus and G. cuneiformis , were separately derived from G. antetexanus . Comparison with gnathodont faunas described from Great Britain, Europe, and Australia indicates worldwide distribution of this lineage. The apparent homotaxis of Gnathodus illustrates the value that has been placed on conodonts as tools for regional and intercontinental correlation.     

Distribution of novel and known repeated elements of Solanum and application for the identification of somatic hybrids among Solanum species

Species-specific repetitive DNA probes are a useful tool for the molecular identification of somatic hybrids. Therefore, the distribution of three repetitive DNA elements of Solanum was investigated in Solanum wild species, Solanum breeding lines, and in more distantly related species of the genera Lycopersicon, Nicotiana, and Datura. The clone pSCH15, obtained from S. circaeifolium, represents a new 168-bp repetitive element; it shows 73–79% sequence similarity to repetitive elements of S. brevidens and Lycopersicon species. The 163-bp element in pSBH6, cloned from S. bulbocastanum, turned out to be very similar (95% sequence homology) to the Lycopersicon element pLEG15/TGRI previously regarded to be present only in species of the genus Lycopersicon and in S. lycopersicoides. Lower sequence similarity of approximately 80% was observed to repetitive elements of S. brevidens which are organized differently. The repeats exhibited different degrees of specificity: by Southern hybridization the element represented by the clone pSBH6 could be detected in almost all Solanum species investigated here but only after long exposure to X-ray film. The previously described Solanum-specific element represented by the clone pSA287 was also found, although in a very low copy number, in Lycopersicon esculentum. Therefore, detection of the repetitive elements pSA287 and pSBH6 in those species in which the respective repeat is less represented depends on exposure time. In contrast, the element pSCH15 is prominently present only in a small number of Solanum wild species and — to some extent — in the diploid breeding lines as revealed after long exposure. Use of these repeated elements for the identification of specific genomes in protoplast-fusion hybrids between Solanum wild species and Solanum breeding lines, or between two breeding lines, was evaluated.     

Frasnian conodonts from the eastern Russian Platform

The Frasnian conodonts of the eastern Russian Platform from the Timan in the north to the Orenburg Region in the south are characterized in detail both paleontologically and stratigraphically. This paper is a continuation of a previously published monograph (Ovnatanova and Kononova, 2001) on the Frasnian conodonts of the shallow-water strata of the central Russian Platform, where polygnathids prevail. The conodont assemblages from the shallow-water and basinal sections of the Volga-Ural province and Southern Timan are analyzed. Problems and difficulties associated with the correlation of the shallow-water and deep-water Frasnian sections of the eastern Russian Platform and their correlation with the existing zonal conodont scales are discussed. The correlation between the deep-water Mendym deposits and the shallow-water Rechitza and Voronezh strata of the Volga-Ural province with the Vetlasyan and Sirachoi deposits of the Southern Timan is substantiated. The stratotype of the Semiluki Regional Stage of the Central Devonian Field contains equivalents of the lowermost beds of the Domanik Formation (unit 1), based entirely on polygnathids. Phylogenetic reconstructions for Palmatolepis and Polygnathus are suggested based on the ontogenetic series for some species of these two genera and the presence of transitional forms between some of the species. Based on these phylogenetic reconstructions, conodont zonal scales for the shallow-water and deep-water sections of the Frasnian Stage of the Russian Platform are proposed and their correlation with the existing conodont zonal scales is also adduced. Some aspects of biofacies control are considered based on the distribution of conodonts in the sections studied. In Systematic Paleontology, 91 conodont species of the genera Ancyrodella, Mesotaxis, Palmatolepis, and Polygnathus are described, including the new species Palmatolepis menneri, P. kaledai, P. acutangularis, and Polygnathus reitlingerae. DEDICATION To the blessed memory of the outstanding geologist Vladimir Vladimirovich Menner     

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.