New Pennsylvanian (Moscovian) echinoderms from Xinjiang Uyghur Autonomous Region,western China

The few Pennsylvanian echinoderm faunas reported from the Paleotethys are from China, Japan, and Australia. The Japanese and Chinese faunas contain camerates that are rare in Pennsylvanian faunas worldwide.Genera of one monobathrid camerate, two disparids, five cladids, one flexible, one blastoid, and one archaeocidarid are reported from the late Moscovian uppermost part of the Qijiagou Formation or lowermost part of the Aoertu Formation in the Taoshigo Valley near Taoshuyuan Village, Xinjiang Uyghur Autonomous Region, China. They are here referred to as the Aoertu fauna, which is most closely allied with Pennsylvanian crinoids reported from a slightly older fauna from the Qijiagou Formation in the Taoshigo Valley and with faunas known from Japan and North America. Most taxa are identified to the genus level and left in open nomenclature because of poor or partial preservation. In addition, camerate ossicles and a cup of Synbathocrinus are reported from the early Moscovian part of the Sanquanzi Formation at Yamansu, southeast of the Taoshigo Valley. New taxa described are: Synbathocrinus labrus n. sp., Stellarocrinus qijiagouensis n. sp. and Metaperimestocrinus aoertuensis n. sp.     

The youngest record of Carcharopsis (Chondrichthyes) from the Pennsylvanian of Xinjiang Uyghur Autonomous Region,China

A chondrichthyan tooth having a serrated edge, from the Late Pennsylvanian of Xinjiang Uyghur Autonomous Region, China, was identified as Edestus sp. [Cheng, Z., Lucas, S.G., Zidek, J., 1996. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1996 (11), 701-707]. Because of the lack of bilateral symmetry, the tooth does not belong to Edestus or any other known edestoid. It is too incomplete for a definite identification, but its features are consistent with Carcharopsis, which also has serrated teeth, and not with any other described late Paleozoic chondrichthyan genus. If it is Carcharopsis, it is the latest occurrence of the genus, which was previously known only from the Mississippian and the Early Pennsylvanian.     

Arachnids from the Carboniferous of Russia and Ukraine,and the Permian of Kazakhstan

New finds of Late Palaeozoic arachnids, based on three well-preserved carapaces from the Carboniferous of Russia and Ukraine and one complete, albeit poorly preserved, specimen from the Permian of Kazakhstan, are described. The spider genus Arthrolycosa is reported from the Late Carboniferous (Late Pennsylvanian: Kasimovian–Gzhelian) of Chunya in the Tunguska Basin of Siberia; it is the first find of a spider outside the Carboniferous tropics. Another fossil assigned to the same genus comes from the Late Carboniferous (Early Pennsylvanian: Bashkirian) of Kamensk–Shakhtinsky in the Donets Basin of Russia; it is probably the oldest fossil spider known. A thelyphonid (whip scorpion) carapace is described from the Late Carboniferous (Late Pennsylvanian: Kasimovian) of the adjacent Lugansk Province of the Donets Basin of Ukraine.     

Pennsylvanian (Late Carboniferous) calcareous microfossils from Cedro Peak (New Mexico,USA). Part 2: Smaller foraminifers and fusulinids

The Pennsylvanian stratigraphic section of the Manzanita Mountains (central New Mexico, USA) is restudied at Cedro Peak. This Pennsylvanian succession is divided into the Sandia Formation, Gray Mesa Fm, Atrasado Fm, and lower part of Bursum Fm. The sampled limestones of the Gray Mesa and Atrasado formations yielded three age-distinctive fossil assemblages: they are of (a) latest Atokan/early Desmoinesian, (b) late early Desmoinesian, and (c) middle Virgilian. The following new taxa of foraminifers are proposed: Endoteboidea n. superfam.; Spireitlinidae n. fam.; Millerellinae n. subfam.; Pseudonovella marshalli n. sp.; Pseudonovella ohioicus nom. nov.; Pseudoacutella n. gen.; Pseudoacutella hoarei nom. nov.; Plectofusulina manzanensis n. sp. Translated or emended names are: Staffelloidea, Ozawainelloidea, Ozawainellidae, and Profusulinellidae.     

THE VEGETATIVE ANATOMY OF SCHOPFIASTRUM DECUSSATUM FROM THE MIDDLE PENNSYLVANIAN OF THE ILLINOIS BASIN

Schopfiastrum decussatum, a monostelic pteridosperm, has been recovered from Middle Pennsylvanian age coal balls from six localities within the Illinois Basin. Additional features of stem anatomy include the presence of horizontal sclerotic plates in the inner cortex, secretory ducts in the inner and outer cortex, and adventitious roots. The primary xylem is interpreted as bilobed, emitting massive leaf traces in a distichous manner. Five orders of frond members are described, including pinnules. All orders of the frond contain secretory ducts and tissues characteristic of Schopfiastrum stems. The rectangular, bilobed protostele, and method of leaf trace origin indicate that Schopfiastrum is more closely related to certain lower Carboniferous seed ferns than to contemporary Pennsylvanian pteridosperms.     

Late Carboniferous palynostratigraphy of the Itararé Subgroup, northeastern Paraná Basin, Brazil

A new biozonation is proposed for the Late Carboniferous strata of the northeastern Paraná Basin (São Paulo and Paraná States), based on well-preserved palynological assemblages recovered from outcrops and core samples retrieved from the Itararé Subgroup. Assemblages include forty-nine species of trilete spores and twenty-five pollen species. The oldest biozone, the Ahrensisporites cristatus Interval Zone (AcZ), in the basal portion of the Itararé Subgroup, includes eleven exclusive spore species, and is assigned a Pennsylvanian (late Bashkirian to Kasimovian) age. The overlying proposed biozone, the Crucisaccites monoletus Interval Zone (CmZ), dated late Pennsylvanian (Kasimovian to Gzhelian), ranges approximately from the top of the lower portion to the middle portion of the Itararé Subgroup, bears only one exclusive species of pollen. Both zones are characterized by the dominance of trilete spores and monosaccate pollen grains, including also few taeniate pollen grains. Furthermore, the basal Protohaploxypinus goraiensis Subzone of the Vittatina costabilis Interval Zone (VcZ) is broadly recognized for the first time in the upper portion of the Itararé Subgroup in the northeastern Paraná Basin. This subzone is attributed to the Early Permian (Early Cisuralian), and is characterized by the dominance of monosaccate pollen grains, a significant increase of taeniate grains (mainly Protohaploxypinus spp.), apart from the appearance of polyplicate pollen (Vittatina spp.). Reference sections and the main characteristics of the two new zones (AcZ and CmZ) are presented, along with a preliminary palynological correlation and discussion on their ages.     

The Carboniferous floras of the Iberian Peninsula: A synthesis with geological connotations

A general review is presented of the Carboniferous floral records in the Iberian Peninsula in the context of the geological history and distribution of the different basins. Mississippian floras are found in Sierra Morena, where major strike-slip faults brought in terranes of diverse provenance. Lower Pennsylvanian floras are represented in the Peñarroya-Belmez-Espiel (Córdoba) and Villanueva del Río y Minas (Sevilla) coalfields of SW Spain (also strike-slip controlled), at La Camocha, near Gijón (Asturias), and in other parts of northern Spain. Middle Pennsylvanian is represented near Oporto, but more completely in the Central Asturian Coalfield, as well as other, more limited localities in NW Spain and the Pyrenees. Upper Pennsylvanian (Stephanian Stage-Cantabrian to Stephanian B substages) floras are splendidly represented in NW Spain. Uppermost Pennsylvanian (Stephanian C-Autunian) floras are present in the strike-slip controlled Douro and Buçaco basins of North Portugal, the Pyrenees, Central Spain (Ciudad Real, Guadalajara, Zaragoza), and SW Spain (Guadalcanal and Valdeviar in Sevilla province).A complete succession of megafloral zones is presented. This includes a new Annularia spicata Zone at the top of the Pennsylvanian (equivalent to middle to upper Autunian which has been often attributed to the Lower Permian). The information is summarised in charts compiled from a selection of the most significant species. A number of floral elements are illustrated including the zonal indices. Brief taxonomic comments are provided in the Appendix.     

THE ULTRASTRUCTURE OF PALEOZOIC FERN SPORES: I. BOTRYOPTERIS

The spores of four species of the Paleozoic filicalean fern Botryopteris are examined at the ultrastructural level. Spores of B. cratis, B. forensis, B. globosa, and an unnamed species from the Lower Pennsylvanian, are compared on the basis of sporoderm stratification and the presence or absence of a sculptine layer. The species examined differ widely as to the type of reproductive unit in which they are borne and include forms that range throughout the Pennsylvanian. In all species the exine is homogeneous, lacking cavities and lamellae. A thin nexine is present in the Middle and Upper Pennsylvanian taxa, but is absent in the Lower Pennsylvanian spores. Only one spore type (B. cratis) possesses a clearly defined sculptine layer. Features of the sporoderm are compared with those of extant, homosporous pteridophyte spores.     

Discovery of radiolaria and conodonts in the Carboniferous–Permian of San Salvador Patlanoaya (Puebla,Mexico); biostratigraphic implications

New identifications of radiolaria and conodonts allow clarifying the biostratigraphy of the Pennsylvanian and Permian deposits in San Salvador Patlanoaya (Puebla State, Mexico). The radiolaria are sometimes relatively common in the series, but weakly diversified and endemic. Among the conodonts, Streptognathodus bellus is characteristic of the Late ‘Virgilian’ sensu Baars, probably coeval with the Early Wolfcampian sensu Thompson or Wilde, the Bursumian of Ross & Ross or the Orenburgian–Asselian stage or substage of Russia (sensu Davydov). San Salvador Patlanoaya is a key-section for the Pennsylvanian–Permian boundary in Mexico.     

The fern Stauropteris oldhamia Binney: New data on branch development and adaptive significance of the hypodermal aerenchyma

Well-preserved specimens of Stauropteris oldhamia are described. The material was collected in the early 1920s from the Lower Westphalian (Early Pennsylvanian) Saurue seam from Belgium. The fossil plants occur as permineralized axes fragments within a coal ball. This study confirms most of the interpretations made by previous researchers. The observation of immature axis however suggests a less regular organization than previously interpreted beyond the three first branching orders. We also highlight the presence of profusely and dichotomously branched aphlebiae, the lack of laminate organs as well as the presence of hypodermal aerenchyma in all plant parts. We interpret these features as part of a very specialized assimilatory apparatus indicating an adaptation to a humid swamp environment.     

Fungal endophytes in Astromyelon-type (Sphenophyta,Equisetales, Calamitaceae) roots from the Upper Pennsylvanian of France

A distinctive fungal endophyte, Cashhickia acuminata nov. gen. et sp., is described from permineralized calamite roots from the Upper Pennsylvanian Grand-Croix cherts of France. Heavily infected roots contain numerous intracellular hyphae in the outer cortex that arise from a meshwork-like mycelium extending between cortical cells. All intracellular hyphae are oriented toward the root center; none occur on the inner periclinal host cell walls. Other roots of the same type show localized infection by this fungus in which isolated cortical cells contain or give rise to intracellular fungal growth. Within the cortical cells are host responses in the form of callosities that indicate the roots were alive at the time of infection. Other endophytes are present in the same host tissue but are less frequent. The discovery of this association provides the first detailed account on the morphology of a Carboniferous fungal root endophyte, as well as the spatial distribution within the host, and infection pathways within the cortical tissues.     

Mississippian (Lower Carboniferous) microfossils from the Chios Mélange (Chios Island, Greece)

The Chios Mélange is a thick Paleozoic wildflysch sequence that crops out on the Greek island Chios. It is composed of chert, limestone and volcanic blocks floating in a siliciclastic turbiditic matrix. New data suggest that the youngest blocks within the Chios Mélange are clasts of a breccia from the Kouramia-Nenitouria area that contain conodonts of late Visean or early Serpukovian age. The conodont fauna from the breccias is characterized by the genera Gnathodus and Lochriea, which favored deeper-water, open-marine facies. Elsewhere on the island, in the Papalia-Nagos area, calcareous microfossils of middle to late Visean age have been recovered from lime grainstone beds within the Chios Mélange. Allochems making up the grainstones are interpreted to have been transported into deeper-water turbiditic facies, but there is no evidence of stratigraphic reworking. Age-diagnostic calcareous microfossils include the alga Koninckopora inflata and foraminifers in the genus Paraarchaediscus. In situ Mississippian microfossils indicate that the Chios Mélange is older than Pennsylvanian (Upper Carboniferous) - Permian, as previously thought. The revised age suggests that the origin of the Chios Mélange may be related to the development of an accretionary prism during the Hercynian Orogeny.© 2003 Éditions scientifiques et médicales Elsevier SAS. All rights reserved.     

Carbon and oxygen isotope evidence for high-frequency (104–105 yr) and My-scale glacio-eustasy in Middle Pennsylvanian cyclic carbonates (Gray Mesa Formation), central New Mexico

We combine cyclo- and sequence stratigraphy along with whole rock δ¹³C and conodont apatite δ¹⁸O analysis to document high-frequency (10⁴–10⁵ yr) and My-scale sea-level changes for the Middle Pennsylvanian (Desmoinesian or Moscovian) Gray Mesa Formation of central New Mexico. Approximately 75 subtidal cycles (1–8 m) are grouped into 4 1/2 My-scale depositional sequences (40–80 m). About 50% of the cycles show evidence of prolonged subaerial exposure at cycle tops with the development of calcretes, diagenetic mottling, and regolith intraclasts. High-resolution δ¹³C analysis of whole rock limestones across nine of the cycles indicates that the cycle tops were diagenetically altered by isotopically light, meteoric fluids during sea-level fall and lowstand. These δ¹³C trends support the interpretation that high-frequency sea-level changes were responsible for cycle development.Conodont apatite δ¹⁸O values from sampled cycles indicate that the high-frequency sea-level changes were driven by glacio-eustasy combined with changes in surface seawater temperature (SST). δ¹⁸O values from conodont apatite, spanning parts of three depositional sequences indicate that My-scale glacio-eustasy and/or SST changes controlled sequence development. δ¹⁸O shifts indicate that the magnitudes of 10⁴–10⁵ yr glacio-eustasy were between ～ 55 and 170⁺ m combined with tropical SST changes of ～ 1.5°–6 °C. Calculated My-scale glacio-eustatic oscillations were between ～ 60 and 140 m with SST changes of < 3.5 °C. The most plausible driver for the My-scale paleoclimate changes is long-period obliquity (～ 1.2 My) variations. These calculated high-frequency, glacio-eustatic values are similar or greater than Pleistocene values, and lie within the range estimated for other Middle Pennsylvanian successions using a variety of independent eustatic proxies. The similarity in range of magnitudes between high-frequency and My-scale sea-level changes combined with the large differences in magnitudes between individual high-frequency sea-level oscillations helps explain the lack of systematic cycle-stacking patterns within these Pennsylvanian icehouse sequences.     

Galtierella biscalithecae nov. gen. et sp., a Late Pennsylvanian endophytic water mold (Peronosporomycetes) from France

A new fossil water mold (Peronosporomycetes), Galtierella biscalithecae nov. gen. et sp., consisting of coenocytic hyphae occurs as an intracellular endophyte in a partially degraded specimen of the reproductive organ Biscalitheca cf. musata (Zygopteridales) from the Upper Pennsylvanian Grand-Croix cherts (Saint-Étienne Basin, France). Some hyphal tips produce small spheres that subsequently develop into ornamented, opaque-walled oogonia; amphigynous antheridia encircle the necks of several immature oogonia. Also present are ovoid structures, which may represent differently shaped oogonia, hyphal swellings, or zoosporangia. Small dome-shaped structures, probably zoospore cysts, are attached to numerous host cell walls. This discovery sheds new light on the morphology and biology of water molds in a Carboniferous ecosystem.     

Large Pelycosaur Footprints from the Lower Pennsylvanian of Alabama,USA

Relatively large tetrapod footprints from Lower Pennsylvanian strata of the Pottsville Formation in the Warrior basin coal field of Alabama were previously assigned to the ichnogenus Attenosaurus. However, three morphotypes are present, and we assign these tracks to Attenosaurus, a new ichnogenus (Alabamasauripus) and Dimetropus. Most of these tracks apparently were made by large pelycosaurs, for which there is no bone record older than Late Pennsylvanian.     

Miofloral succession and interpretation of the base of the permian system in the Eastern Shelf of North Central Texas,U.S.A.

The study includes palynological investigation of the Pennsylvanian (Upper Carboniferous) and lowermost Permian (Wolfcampian) strata of Texas, with special emphasis on the Pennsylvanian and Permian systemic boundary in the Eastern Shelf of north central Texas. The following palynozones have been recognized: (1) Grumososporites-Simozonotriletes Zone (Atokan); (2) Torispora-Thymospora Zone (Demoinesian); (3) Cristatisporites—Cadiospora arcuata Zone (Missourian); (4) Crucisaccites americanus Zone (Early Virgilian); (5) Guptaesporites honakerensis Zone (Late Virgilian); and (6) Nuskoisporites crenulatus-Hamiapollenites saccatus Zone (Early Wolfcampian). The base of the Permian Wolfcampian Stage has been placed at the base of Nuskoisporites crenulatus-Hamiapollenites saccatus Zone, which approximately coincides with the base of the Coleman Junction Limestone Member of the Putnam Formation of the Wichita Group. This conclusion may appear to be contradictory to fusulinid evidence because it places the base of the Permian System 182.88 m above the so-called fusulinid boundary. It also disagrees with the paleobotanical boundary. The discussion concludes that fusulinid and paleobotanical evidence is inconclusive.     

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.     

The ultrastructure and reproductive significance of Lasiostrobus microspores

The normally inaperturate microspores of the Pennsylvanian cone Lasiostrobus are found to rarely possess trilete sutures, confirming the subequatorial position of the sacci. Sporoderm ultrastructure compares favourably with saccate pollen of some extant members of the Coniferales. The shedding of grains in the microspore stage suggests that microgametophyte evolution had passed the stage of prothallial cell production, and that Lasiostrobus is an advanced member of the Paleozoic gymnosperm complex.     

A new species of Dyoros (Brachiopoda; Chonetoidea) from Nevada (United States) and stratigraphic implications for the Pennsylvanian and Permian Antler Overlap assemblage

Newly discovered fossil localities in coarse-grained deposits of the Pennsylvanian and Permian Antler overlap assemblage in the southern Shoshone Range, north-central Nevada have yielded a low-diversity assemblage consisting chiefly of a new species of chonetoidean brachiopod: Dyoros (Lissosia) nevadaensis nov. sp. The subgenus Dyoros (Lissosia), is known from Leonardian and lower Guadalupian strata in North America, mainly in Texas.The coarse-grained lithology of the host strata, their unconformable relation on deformed lower Paleozoic rocks, and the Leonardian and(or) lower Guadalupian age indicated by Dyoros (Lissosia) provide evidence that host strata are younger than strata of the Antler overlap assemblage in nearby areas of the southern Shoshone Range and suggest that an unconformity of local extent may be present within the overlap assemblage. The fossil age ranges and lithologic data suggest that the host strata may be correlative with the Guadalupian Edna Mountain Formation, an unconformity-bounded unit that forms the upper part of the Antler sequence in the Battle Mountain area to the north. This correlation suggests that the unconformity beneath these strata may have regional extent in north-central Nevada. The origin of the inferred regional unconformity is unknown and may have resulted from relative changes of sea level or regional extensional or contractional tectonism in the area of the former Antler highlands, which forms the substrate for the Antler overlap assemblage.     

STUDIES OF PALEOZOIC MARATTIALEANS: THE MORPHOLOGY AND PHYLOGENETIC POSITION OF EOANGIOPTERIS GOODII SP. N.

Eoangiopteris goodii sp. n. is described from Upper Pennsylvanian coal balls from Ohio (Shade locality) containing isolated pecopterid pinnules approx. 7 × 9 mm that bear up to 20 linear synangia on the lower surface. The synangia extend at right angles from the midrib to the downturned pinnule margins and measure 2.0–3.5 mm in length. Individual synangia are compact and are composed of 10–19 sporangia that have their bases embedded in an elongate parenchymatous pad. In longitudinal section sporangia measure 0.4 × 2.0 mm and have acute elongate, curved apices. Spores average 70 μm in diam and are most similar to the dispersed spore genus Verrucosisporites. Eoangiopteris goodii differs from the generitype E. andrewsii Mamay in its greater size, pinnule histology, and spore type. Sporangium wall complexity and spore type of the two presently known species of Eoangiopteris are considered to be at about the same evolutionary level as the more primitive species of Scolecopteris. Construction of the synangia in Eoangiopteris is different from that of Scolecopteris and indicates that at least two evolutionary lines are recognizable within the Pennsylvanian Marattiales.