LYCOSPORA FROM CARBONIFEROUS LEPIDOSTROBUS COMPRESSIONS

Spores were extracted from Carboniferous Lepidostrobus compressions in order to associate in situ microspores with dispersed species of Lycospora. Two hundred twenty-six cones were examined, of which 61 contained spores. Fertile cones came from the Westphalian D of England, Namurian B through Westphalian D of the Appalachian and Illinois basins, and the Westphalian D of the Western Interior. Cones were separated into species based on microspore and cone morphology. Lycospora trigonoreticulata was produced by Lepidostrobus princeps from Westphalian C-D rocks from Missouri, the Illinois Basin, and the Appalachian Basin. Lycospora rotunda was produced by Lepidostrobus sp. A from Westphalian A rocks of Alabama. Two cone species produced Lycospora torquifer: Lepidostrobus praelongus from the Westphalian D of Pennsylvania and Lepidostrobus variabilis from the Westphalian A and C of the Illinois and Appalachian basins. Lycospora punctata was produced by Lepidostrobus cf. squarrosus from the Westphalian D of England, the Appalachian Basin, and Illinois Basin. Lycospora noctuina was produced by Lepidostrobus haslingdenensis from the Namurian B/C of Illinois. Microspore species are differentiated primarily on the basis of size, cingulum structure and width, and ornamentation. Cone species differ in width and distal lamina size, shape, and attitude. Lycospora species isolated from clastic species of Lepidostrobus differ completely from those of coal-swamp species, confirming that lycopod trees from clastic environments represent biologically different species from those centered in coal swamps.     

SOURCE PLANTS FOR CARBONIFEROUS MICROSPORES: LYCOSPORA FROM PERMINERALIZED LEPIDOSTROBUS

Permineralized Lepidostrobus specimens were examined from the Westphalian A of England, Westphalian B (Breathitt) of the Appalachian Basin, and the Westphalian D (Desmoinesian) of the Illinois Basin and Western Interior Coal Region to correlate species of the dispersed spore genus Lycospora with source cones and cone types (“species”) with parent plants. Based on 51 specimens of Lepidostrobus oldhamius sensu Balbach, three cone types, producing three species of Lycospora, are recognized. Lycospora pellucida occurs in smaller diameter cones with a prominently coronate stele; cones are associated with Lepidophloios harcourtii from the Westphalian A-B. Lycospora pusilla occurs in larger diameter cones with a small “pith” containing elongate, sclerified, tanniferous cells from Westphalian D coal balls. Secretory cells also are present in sporophyll trace sheaths and in subarchesporial parenchyma pads in sporangia. The suggested plant source is Lepidodendron hickii. Lycospora granulata occurs in large cones with a large “pith” composed of thin-walled parenchyma; no secretory cells were observed in the “pith,” sporophyll traces, or subarchesporial pads. These cones are associated with Lepidophloios hallii from the Westphalian D. Association of Lycospora species with source lycopods increases the possibilities of paleoecological interpretation using microspores from Carboniferous coals.     

Stratigraphic occurrences and vegetational patterns of Pennsylvanian pteridosperms in Euramerican coal swamps

The vegetation of many Euramerican coal swamps is known from coal-ball peats in the tropical rainy zone across the paleocontinent of Laurussia in the Upper Carboniferous. The stratigraphic occurrences of coal balls in Europe (lower Westphalian) and in North America (upper Westphalian and lower Stephanian) are largely complementary and a revised correlation chart is given. Quantitative analyses of the vegetation from these autochthonous peats are combined with coal palynology to determine on overview of swamp vegetation patterns during the Westphalian and early Stephanian. In comparison to the dominance and great diversity of pteridosperms in the Upper Carboniferous compression floras, seed ferns were minor to subdominant elements in the coal-ball peats. The quantitative composition of organ assemblages of the principal pteridosperms is given with stratigraphic occurrences of Medullosa and Sutcliffia (Medullosaceae), Heterangium, Lyginopteris, Microspermopteris, and Schopfiastrum (Lyginopteridaceaea) and Callistophyton (Callistophytaceae). The major peat contributors among pteridosperms are Lyginopteris (?90%) in the Westphalian A and Medullosa (?95%) in the Westphalian D and Stephanian. Lyginopteris became extinct in the early Westphalian B and Medullosa became abundant during the Westphalian C–D transition and a subdominant in the Stephanian contributing 13–21% of the permineralized peats. Paleoecological interpretations indicate that medullosan trees were most abundant but patchy in distribution in drier swamp stages with enriched nutrients and near major channels in swamps during the Westphalian D. It is suggested that the swamp pteridosperms are probably quite similar to or identical with some of the representatives in the more diverse compression floras. The evolution and diversity noted in swamp pteridosperms are considered principally the product of adjacent lowland communities which repeatedly introduced seeds into the wetlands. The swamp pteridosperms exhibit the greatest fluctuation in abundance (biomass) from site to site in the same coal, probably the highest level of diversity and the only steady pattern of increase in importance during the Late Carboniferous with the possible exception of the Westphalian B–C interval. The close relationships inferred between swamp pteridosperms and their lowland seed sources are also compatible with the frequent association of tree ferns and seed ferns during the Westphalian D and in the Stephanian when these two kinds of frond-bearing trees formed most of the vegetation in both lowland and swamp environments.     

New Euthycarcinoids and an Enigmatic Arthropod from the British Coal Measures

Two new species of euthycarcinoids (Arthropoda), Kottixerxes anglicus sp. nov. and Smithixerxes pustulosus sp. nov., are described from the Coal Measures of Westhoughton, Lancashire and Coseley, West Midlands (Westphalian A and B respectively). Both genera are previously known from Mazon Creek, USA (Westphalian D). An additional, enigmatic arthropod with possible euthycarcinoid affinities, Arthrogyrinus platyurus gen. et sp. nov., is described from Coseley. Hypotheses concerning the phylogenetic position of euthycarcinoids are critically reviewed.     

A New Japygid Dipluran from the Lower Cretaceous of Brazil

The first Mesozoic japygid (Hexapoda: Diplura), Ferrojapyx vivax gen. et sp. nov., is described from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil. There are only two previously described occurrences of japygids in the fossil record: in Miocene or Pliocene onyx marble from Arizona and from the Carboniferous (Westphalian D) Francis Creek Shale of Mazon Creek, Illinois.     

First Mesozoic Scutigeromorph Centipede, from the Lower Cretaceous of Brazil

The first Mesozoic scutigeromorph centipede (Myriapoda: Chilopoda), Fulmenocursor tenax gen. et sp. nov., is described from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil. Previously described fossil Scutigeromorpha are known from Dominican and Baltic amber, the Carboniferous (Westphalian D) Francis Creek Shale of Mazon Creek, Illinois, the Silurian and Devonian of Britain, and the Devonian of New York State.     

A comparison of Asturian lepidodendroid lycophytes from the USA and the Euramerican Variscan Front

Species belonging to the Carboniferous genera Lepidodendron and Lepidophloios are well known from the coalfields of Pennsylvania (Lower Asturian (Westphalian D) substage of the Pennsylvanian) through the publications of Lesquereux and White. They believed that the lycophyte flora was rather distinct from other contemporaneous coal floras elsewhere due to its isolation. A full revision of the lepidodendroid lycophyte species is given here and this reassessment permitted comparisons with those floras known from the European and Canadian Maritimes coalfields. Differences in species distribution suggest there was a barrier (Arcadian Highlands) to species distribution to and from the coalfields of Pennsylvania. Several species are recognized only from the coalfields of Pennsylvania, although the number has been significantly reduced from the original publications.     

Palaeogeography of the late palaeozoic in the bohemian massif

The palaeogeographic and palaeoclimatic setting of the Bohemian Massif is documented with maps for the following seven stages: Westphalian A, Westphalian B to Westphalian D, Stephanian A, Stephanian B, Stephanian C to Autunian, Saxonian and Thuringian. The evaluation of present occurrences and of the presumed area of sedimentation makes the palaeogeographic picture more comprehensive. Because of the relatively long time intervals represented on the maps, the palaeogeography is expressed in terms of the ratio of different environments in the total sedimentary fill. For the units deposited under a predominantly humid climate, we distinguish between sediments of foothills, alluvial fans, flood plains, lacustrine deltas, perennial lakes, peats and swamps. For the units deposited under semi-arid to arid climatic conditions, sediments of pediplains, alluvial fans or of the alluvial apron were suggested for the margins, and sediments of flood plains, ephemeral and perenial lakes, peats and swamps for the central part of the sedimentary basins. The development of the sedimentary area and of its fill was influenced by diastrophism and particularly by the climate. The rate of sedimentation was controlled by the contemporaneous subsidence. Changing climatic conditions exerted a strong influence on the regionally important fluctuations in the size and shape of the area of sedimentation and on the character of the material deposited. From Westphalian to Austinian times the area of sedimentation was generally expanding. During the Saxonian and Thuringian, the area of sedimentation was reduced. This reduction was caused by the tectonic stabilization and peneplanation of the central part of the Bohemian Massif and the appearance of typical desert climatic conditions.     

Dicksonites pluckenetii(Schlotheim) Sterzel and its affinity with Callistophytales

Female and male (?) fronds of the Stephanian Dicksonites pluckenetii(Schl.) Sterz. initially described by Grand'Eury (1905) have been reinvestigated. Seeds were attached abaxially and subapically to strongly reduced linear pinnules of pinnae which are otherwise similar to sterile ones. Associated synangia are of the Pterispermostrobus type. They appear to have occupied marginal or submarginal position on modified pinnules of irregular outlines. The affiliation of D. pluckenetii with the order Callistophytales is supported. Considering that the oldest (Westphalian) members of D. pluckenetii had unmodified seed-bearing fronds, the progressive reduction of fertile fronds in the evolution of the Callistophytales is suggested. Further stages of the process are observed in the Permian family Trichopityaceae (order Peltaspermales). In another lineage of the Callistophytales the fertile fronds were planated and leafless.     

Neuropteris attenuata, a narrow-stemmed, leaning or lianescent seed fern from the Upper Pennsylvanian of Lower Saxony, Germany

A stem segment and frond of Neuropteris attenuata Lindley et W. Hutton (Pteridospermopsida, Medullosales) are described from the lower Westphalian D (Upper Pennsylvanian) of the Piesberg quarry near Wallenhorst (north of Osnabrück, Lower Saxony, northwestern Germany). The narrow diameter of the stem (3–4 cm), conspicuously long internodes (>20 cm), and the large size of the frond (90 cm long) strongly suggest that N. attenuata was a non-self-supporting, leaning or lianescent plant.     

Phylogénie générale des Calliptéridiacées (Pteridospermopsida)

The study of the evolutionnary trend of the leaf architecture in the genus CallipteridiumWeiss, from the upper Westphalian and Stephanian, has allowed to determine, by extrapolation, the characters that had to show the frond of the direct ascendants which belong to the genus MargaritopterisGothan, from the Westphalian. This clue has led the quest on the field and has enabled the discovery of data which verify the correctness of the formulated hypothesis. This positive result and the evolutionnary trend ascertained by this new portion of the phylum led us to propose a new hypothesis about the characters of the fronds of the more ancient representatives of the phylum, that one can trace with great probability up to the lower Namurian and more hypothetically up to the Dinantian.This work has carried otherwise a revision of the corresponding species from the North of France and Sarre-Lorraine coal basins with a certain number of consequences about stratigraphical correlations.     

Systematics of the Late Carboniferous Medullosalean Pteridosperm Laveineopteris and Its Associated Cyclopteris Leaves

Laveineopteris is emended to become a morphogenus for the vegetative parts of certain medullosalean pteridosperms that grew in the Westphalian (Late Carboniferous) tropical rainforests. The plant bore pinnate foliage as sun leaves and entire orbicular leaves (cyclopterids) as shade leaves. The cyclopterids were attached to the axis of the saplings and epiphyllously to the lower parts of the pinnate fronds. Species recognition is mainly by the macro–morphology of the pinnate fronds. However, species can also be distinguished based on the epidermal anatomy of both pinnate fronds and cyclopterids, in particular the distribution and detailed structure of the stomata and hydathodes. Laveineopteris first appeared in the lowland swamp forests of Europe during the early Langsettian (Westphalian A), and there were two principal species: L. loshii and L. tenuifolia . During the early Bolsovian (early Westphalian C), a variety of L. tenuifolia appeared that was adapted to both the lowland swamps, and the upland swamps of the intramontane basins (central Bohemia and Saar–Lorraine). The smaller–pinnuled L. rarinervis also first appeared in the Bolsovian and seems to have been restricted to the lowland swamps. The morphogenus Cyclopteris should be used only for cyclopterid fossils, where they cannot be related to the species of Laveineopteris , Callipteridium or Margaritopteris that originally bore them.     

A new late Westphalian fossil marattialean fern from Nova Scotia

Sydneia manleyi gen. et sp. nov. is based on part of a fertile frond from the upper Westphalian D of the Sydney Coalfield, Nova Scotia, Canada. It has small synangia composed of laterally fused sporangia that are elongate and with a circular cross-section. The sporangia yielded variably sized monolete and trilete spores with laevigate and microspinate ornamentation; intermediate forms were also observed. The spores can be correlated with the sporae dispersae species Latosporites minutus , Punctatosporites oculus and Laevigatosporites minimus . Size distribution of the spores is variable and highly skewed, suggesting heterogeneity of the spores within the sporangium. Spore ultrastructure indicates that the fossil is part of a fern, and the morphology of the spores and synangia indicate marattialean affinities.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 199–212.     

The age of the British upper coal measures with reference to their miospore content

Details are given of miospore distributions and zonation in the British Upper Coal Measures. These are compared with miospore distributions in strata of comparable age in France, Belgium, The Netherlands, Germany, Spain, and parts of Russia and the United States. Various published correlations of these strata are discussed and an amended correlation between the British and French sequences suggested. It is concluded that there is as yet little palynological evidence for the presence in Britain of Carboniferous strata of an age younger than that of the Westphalian D stage.     

Ein neuer Insektenfund aus dem Westfalium von Ibbenbüren (Westdeutschland)

From rocks of Westphalian D age of Ibbenbueren in Westphalia/W-Germany a new taxon of the Omaliidae Handlirsch, 1906 -Kelleropteron kaelberbergense n. g., n. sp. — is described. It is represented by only one wing. The new genus is characterized by the following features: the subcosta ends on the costa; the radius is very prominent and runs far distantly from the anterior border of the wing proximally; the branches of the sector radii and the medialis are directed distally, not posteriorly; a connective vein between medialis and sector radii is present; the cross-veins are relatively dense and irregularly branched. The Omaliidae are redefined, the relationships of their genera are discussed. The genera are arranged in 3 groups (Fig. 1). In theHeterologies-group - as in the Paoliidae — the subcosta ends on the radius; in theCacurgus — group the subcosta ends on the costa, a connective vein between medialis and sector radii is not present; in theOmalia- group the subcosta also ends on the costa, but a connective vein between medialis and sector radii has evolved.Kelleropteron is a member of the last group, differing fromOmalia above all by its denser, more irregularly arranged cross-veins. The Omaliidae are confined stratigraphically to the Westphalian and geographically to the EuropeanNorthamerican region. They apparently are descendants of the Paoliidae Handlirsch, 1906. In both families a connective vein between medialis and cubitus is present, and the anterior branch of the medialis is lost. The most important difference between them is seen in the cross-veins: within the Paoliidae they are very dense, similar to the archedictyon of the Palaeodictyoptera, but within the Omaliidae they are more widemeshed and more prominent.     

Verkieste Amöben in Steinkohlen aus dem Ruhrgebiet — erster Nachweis vonArcella Ehrenberg im Paläozoikum

Within bituminous coals of Westphalian B age pyritized amoebae of the order Testacea (thecamoebians) were detected. They may belong to the genusArcella. The appearance of this genus, known until now only since the Pleistocene, is extended far into the Palaeozoic by this observation.     

Two species of Selaginella cones and their spores from the Bohemian Carboniferous continental basins of the Czech Republic

Two species of Selaginella cones from the Bohemian Upper Carboniferous continental basins of the Bolsovian and Westphalian D age are described, together with their in situ spores. Two specimens of Selaginella gutbierii yielded microspores closely comparable with the dispersed species Cirratriradites saturnii and megaspores closely comparable with the dispersed species Triangulatisporites vermiculatus. Microspores closely comparable with the dispersed species Cirratriradites annulatus and megaspores resemble the dispersed species Triangulatisporites tertius were isolated from cones of Selaginella cf. leptostachys. All the spores isolated from one cone are of the same type and would be referred to one dispersed micro- and megaspore species if found as Sporae dispersae. The paper contains a review of all palynologically studied Carboniferous Selaginella and Selaginella-like cones and reviews of all in situ and dispersed Cirratriradites and Triangulatisporites spores.     

The Westphalian fossil floras from the Cattybrook Claypit,avon (Great Britain)

Cattybrook Claypit, on the northwest margin ofthe Bristol-Somerset Coalfield, yields an upper Westphalian A (Middle Carboniferous) fossil flora. Most of the fossils are of small plant fragments, transported mainly by winds into floodbasin muds. They represent a variety of plant-types, including pteridosperms, ferns, cordaites, sphenophytes and lycophytes. Of particular interest are finely preserved examples of Lonchopteris, and Corynepteris, which are generally scarce in British floras of this age. Also found at Cattybrook are plant fragments in a crevasse-channel sandstone. These are preserved nearer their original position of growth, and are thus more complete than those found in the floodbasin deposits. They include almost complete fronds of Karinopteris attached to a thick lianatype stem, and large pieces of a Sphenophyllum plant. This is now the best documented late Westphalian A flora in Britain.