CHARLIEA MANZANITANA,N. GEN., N. SP., AND OTHER ENIGMATIC PARALLEL-VEINED FOLIAR FORMS FROM THE UPPER PENNSYLVANIAN OF NEW MEXICO AND TEXAS

Charliea is a new genus (type-species: C. manzanitana), based on pinnately compound leaf material from the richly fossiliferous Virgilian (Upper Pennsylvanian) shales of the Kinney Brick Company quarry near Albuquerque, New Mexico. In several features Charliea resembles Russellites or a zamioid cycad. It has linear-oblong pinnae with broad, oblique attachment and a truncate tip, which is deeply incised to form two to four nearly equal lobes. The venation is simple, parallel, and sparingly dichotomous, each vein ending at the distal margin. The Kinney beds also contain Plagiozamites planchardi, another zamioid form with parallel-veined pinnae, differing from Charliea chiefly in having rounded tips and veins ending in the denticulate margins. An unnamed third form (genus B) in the Kinney beds has long, narrow pinnae with parallel veins and blunt tips; this strongly resembles the Mesozoic conifer Podozamites, but may just as well represent a cycadophyte. Another unnamed taxon (genus A), from an Upper Pennsylvanian deposit in Jack County, Texas, resembles genus B or Russellites in general shape and venation, but the critical distal margins are unknown. In their single-ordered parallel venation, these four foliar types contrast sharply with the two-ordered pinnate venation of most Pennsylvanian fern-like leaves, and seem to foreshadow Mesozoic morphologies. This tendency toward precocious evolution of parallel-veined foliar form in North America is also expressed by a single occurrence of the Asiatic, Permian genus Tingia in the Lower Pennsylvanian of Utah, and by the presence of the predominantly Triassic cycadeoid genus Pterophyllum in the Lower Permian of Texas.     

BOSTONIA PERPLEXA GEN. ET SP. NOV., A CALAMOPITYAN AXIS FROM THE NEW ALBANY SHALE OF KENTUCKY

Bostonia perplexa, gen. et sp. nov., was collected from the Lower Mississippian Falling Run member of the Sanderson Formation. The single short segment of an axis, preserved as a petrifaction, contains at least three vascular columns, each with both primary and secondary tissues. Primary xylem is two or three ribbed, and contains several mesarch protoxylem strands. Gymnospermous secondary xylem is characterized by both uniseriate and multiseriate rays. The ground tissue is parenchymatous except for a few clusters of sclerotic cells. In its apparent polystelic nature, the specimen superficially resembles members of the Pennsylvanian to Permian Medullosaceae. All evidence currently available, however, leads to the conclusion that this species should be placed in the Upper Devonian to Lower Mississippian Calamopityaceae. It has not been determined with certainty whether the species is polystelic (in the sense of the Medullosaceae), or whether the apparent polystely is the result of stelar branching proximal to the level of branch divergence.     

STUDIES OF PALEOZOIC MARATTIALEANS: NEW SPECIES OF SCOLECOPTERIS (MARATTIALES) FROM THE PENNSYLVANIAN OF NORTH AMERICA

Two new species of the late Paleozoic fern Scolecopteris (Marattiales) are described and their relationships within the genus are discussed. Scolecopteris charma sp.n., from Steubenville, Ohio (Duquesne Coal, Upper Pennsylvanian), is similar to species in the Oliveri group, while S. gnoma sp.n. from Providence, Kentucky (Baker Coal, Middle Pennsylvanian), compares favorably with the Latifolia species group. Scolecopteris gnoma is most similar to S. fragilis but differs in its smaller synangia and spore type. S. charma appears generally similar to S. iowensis because of its large pedicel and histologically undifferentiated walls, but differs in a number of characters such as vasculature and spore type. Despite its occurrence late in the Pennsylvanian, S. charma is thought to possess a number of primitive character states (large trilete spores, vascularized pedicels, flat pinnules with downturned margins). Using the same criteria for the much older S. gnoma, we note a number of relatively apomorphic character states (small monolete spores, unvascularized pedicels, extended pinnule margins). An outgroup analysis of species-level characters of Scolecopteris gives a better concept of primitive versus derived traits in marattialean and other ferns. Genera in the Paleozoic fern orders Filicales (Ankyropteris) and Zygopteridales (Corynepteris, Musatea) were chosen as outgroups, and the comparisons support suggestions for the polarity of several important characters. Some of these agree with previously proposed evolutionary polarities based on the geological occurrence of marattialean ferns.     

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.     

QUAESTORA AMPLECTA GEN. ET SP. N., A STRUCTURALLY SIMPLE MEDULLOSAN STEM FROM THE UPPER MISSISSIPPIAN OF ARKANSAS

A well preserved, permineralized seed fern stem is described from the Upper Mississippian Fayetteville Formation of north central Arkansas. Quaestora amplecta gen. et sp. n. is 41.6 cm long and exhibits six pairs of decussate, highly decurrent petiole bases. The stem has a cruciform, exarch protostele with prominent secondary xylem, vascular cambium and secondary phloem. Leaf traces are terete and occur as an outer ring with a small number of internal strands. The cauline vasculature, leaf-trace production, petiolar anatomy and several other features indicate that this specimen represents the most structurally simple and geologically ancient medullosan stem presently recognized.     

ARCHAEOCALAMITES LAZARII,SP. NOV.: THE RANGE OF ARCHAEOCALAMITACEAE EXTENDED FROM THE LOWERMOST PENNSYLVANIAN TO THE MID-LOWER PERMIAN

Archaeocalamites lazarii, sp. nov., is based on a few vegetative compressions from the mid-Lower Permian Leonard Series near Fulda, north-central Texas. It is a minor component of a rich biota that includes sphenophylls, pteridophylls, conifers, conchostracan crustaceans, eurypterids, arachnids, insects, xenocanth sharks, and coprolites attributed to tetrapods. Biota and sedimentology indicate deposition in a small freshwater body on a deltaic floodplain. Leaves of A. lazarii differ in size and posture from those of the widespread and characteristically Mississippian aggregate species A. radiatus; nonpreservation of rooting and reproductive organs prevents interpretation of its phylogenetic relationship with other equisetaleans. The main significance of A. lazarii is that it extends the recorded stratigraphic range of the Archaeocalamitaceae from the lowermost Pennsylvanian (Namurian B) to the mid-Lower Permian (Artinskian), leaving a hiatus in records of approximately 55 Ma. This Lazarus taxon also occurs with possibly the youngest recorded eurypterid.     

PHILLIPOPTERIS GEN. NOV.—ANATOMICALLY PRESERVED SPORANGIAL FRUCTIFICATIONS FROM THE UPPER PENNSYLVANIAN OF THE APPALACHIAN BASIN

The discovery of a new type of sporangial fructification in coal balls from the Upper Pennsylvanian of Ohio provides the basis for describing Phillipopteris globiformis gen. et sp. nov. Sporangia are borne terminally on up to two orders of branching axes. Penultimate axes branch pinnately to produce irregularly branched ultimate axes. Sporangial wall cells are of a single type and show no specialization for dehiscence. Spores are radial and trilete, and reminiscent of the sporae dispersae genus Dictyotriletes. Phillipopteris increases our knowledge of diversity among fernlike plants from the late Paleozoic, and shares several features with Sclerocelyphus Mamay.     

STEM-ROOT TRANSITION OF AN UPPER PENNSYLVANIAN WOODY LYCOPSID

Anatomically preserved specimens of a woody lycopsid showing the transition from the stem to the rooting region are described from the Upper Pennsylvanian Duquesne Coal of Ohio. Specimens have exarch protosteles that are apparently medullated at distal levels and exhibit abundant secondary xylem. Cortical tissues accompanying the stems have periderm, and show leaf bases or cushions. Although features of the stems are compatible with those of the arborescent Lepidodendrales, the plants have a rounded cormose rooting region, rather than the much-branched and elongated stigmarian system usually associated with the order. Specimens of this type expand our knowledge of the diversity among Paleozoic lycopsids and document the occurrence of representatives with an Isoetes-like base in Pennsylvanian strata.     

A new Serpukhovian (Mississippian) fossil flora from western Argentina: Paleoclimatic, paleobiogeographic and stratigraphic implications

An uppermost Mississippian (Serpukhovian) fossil flora from western Argentina is described for the first time. Its particular composition includes both, warm climate plants typical from the Pennsylvanian of Gondwana, namely pteridosperms, sphenophytes as well as arboreous lycopsids; and small lycopsids characteristic of cooler climates of pre-Serpukhovian times. A warm–temperate climate is inferred for this association, which fills a time gap between those Pennsylvanian and Mississippian classic floras previously known. The presence of Tomiodendron, Nothorhacopteris kellaybelenensis and Paracalamites allow the recognition of the Paraca Floral Realm in Argentina, supporting the existence of a nonglacial paleoclimate period, developed just before of the Bashkirian–Moscovian Gondwanan glaciation and after a latest Visean glaciation. It likewise would indicate the paleogeographic position of south west Gondwana, which would have acceded to a warm temperature latitudinal belt during Serpukhovian times. The recognition of this floral assemblage has relevant biostratigraphic and lithostratigraphic implications for the western margin of Gondwana. It defines a new zone for western Argentina, herein named Frenguellia eximia–Nothorhacopteris kellaybelenensis–Cordaicarpus cesarii (FNC) zone, which is recorded in the oldest known lithostratigraphic unit from the Paganzo Basin, the Loma de los Piojos Formation nom. nov.     

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.     

Rugose corals and associated carbonate microfossils from the Brigantian (Mississippian) of Castelsec (Montagne Noire, southern France)

The disused quarry east of Castelsec offers a view of shallow-marine carbonates of the poorly known Uppermost Mississippian of the Montagne Noire. At Castelsec, sections are studied in two characteristic facies types (bioclastic wackestone and microbial dominated boundstone) of the Upper Mississippian. The succession is rich in rugose corals and carbonate microfossils. Six genera with seven species belonging to a rugose coral fauna consisting of at least eight genera with several species are described herein; Dibunophyllum castelsecensis sp. nov. is described as new. Twenty-seven carbonate microfossils of different groups have been identified. The Castelsec succession is Brigantian in age, based on the stratigraphic occurrence of rugose corals, foraminifers, and calcareous algae observed in both sections. The rugose coral fauna shows relationships with the well-known fauna of northwestern Europe and the Ouralian-Asian Province. Typical elements of northwestern Europe are missing at Castelsec and vice versa. This differentiation between north and south is interpreted as responses to different palaeolatitudes and tectonic settings.     

Lacoea,a lower Pennsylvanian noeggerathialian cone from Illinois

Lacoea seriata is described from 2- and 3-dimensional compression fossils from the Lower Pennsylvanian (Westphalian A?-B?) strata of west-central Illinois. Stratigraphic data are given for the fossiliferous shale and siltstone which occur within basal Pennsylvanian fill in a depression within Mississippian Limestone. The six specimens studied include isolated fringed semicircular disks and a cone segment showing the arrangement of the sporophylls on the axis. Comparisons are made with Discinites, Noeggerathiostrobus, Saarodiscites, Heninia and Tongshania. Lacoea exhibits morphological characteristics of the noeggerathiales.     

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.     

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.     

Paleozoic seed fern pollen organs

Paleozoic pollen organs exhibit numerous morphological forms that have been arranged in categories based on their probable organization. Progymnosperm ancestors are characterized by three dimensional branching systems bearing pairs of terminal sporangia. Early Mississippian examples of seed fern fertile branches appear little modified from the progymnosperms. These pteridosperm microsporangia are nonsynangiate and thin walled with longitudinal dehiscence. By Upper Mississippian time all forms show sporangial clustering into large or small groups, with several taxa exhibiting radially symmetrical synangia. In the Lower Pennsylvanian all pollen organs are synangiate and appear to consist of a uniseriate ring of sporangia that either surround a central hollow, or are bilaterally flattened. Sporangial dehiscence in all forms is longitudinal and toward the center of the synangium. In bilateral synangia with no central hollow, the sporangia either separate laterally or effective dehiscence areas are restricted to the free apical portions of the sporangia. Callistophytacean synangia resemble the lyginopterid type, but are abaxial on laminar foliage. This family is thought to have evolved from the lyginopterids during the Early Pennsylvanian. Middle Pennsylvanian medullosan pollen organs are all radial and may be solitary, aggregated into groups, or fused into a large compound synangium. Several pollen organ types are reinterpreted, and the possible evolutionary relationships among the various Paleozoic pollen organ forms are discussed based on synangial organization, patterns of frond branching, and pollen or prepollen morphology.     

MITROSPERMUM VINCULUM SP. NOV., A CARDIOCARPALEAN OVULE FROM THE UPPER PENNSYLVANIAN OF OHIO

Numerous anatomically preserved ovules assignable to the genus Mitrospermum have been discovered in Upper Pennsylvanian sediments of Eastern Ohio. Although basically similar to Mitrospermum compressum, the newly discovered specimens exhibit several consistent differences. Ovules are strongly platyspermic, up to 4.2 mm long, 4.0 mm wide, and 0.6 mm thick. In the minor plane, ovules are broadest at the base and taper toward the micropyle. The integument exhibits three topographic regions: endotesta, sclerotesta, and sarcotesta. The sarcotesta is extremely broad in the major plane, where it forms two membranous wings. A single terete vascular bundle enters the base of the ovule, traverses the integument, and divides to form two integumentary bundles and a conspicuous nucellar platform. Integumentary bundles extend toward the tip of the ovule at the margin of the sarcotesta and sclerotesta. A pollen chamber with a prominent nucellar beak is delimited at the tip of the nucellus. Consistent differences in vascularization, size, nature of the seed base, features of the pollen chamber, and the Late Pennsylvanian age demonstrate that the specimens represent a distinct species. The discovery of these ovules extends the stratigraphic range of Mitrospermum to the Upper Pennsylvanian of Ohio.     

Suavitas imbricata gen. et sp. nov., an anatomically preserved seed analogue of putative lycophyte affinities from upper Pennsylvanian marine deposits

A megasporangiate fructification has been discovered in dysoxic, offshore marine shales from Upper Pennsylvanian deposits of northcentral Texas. This specimen, described as Suavitas imbricata gen. et sp. nov., consists of imbricating, helically arranged sporophylls that diverge from a stout axis. The axis is characterized by parenchymatous pith and weakly developed vascular tissue. Sporangia are terminal with a horizontally elongated, apical dehiscence slit. There is one large functional megaspore in each sporangium, and the megaspore wall is constructed of interconnected rodlets. Parsimony analysis indicates that Suavitas may be related to rhizomorphic lycophytes. The unique combination of characters displayed by this species illustrates the central role played by extinct species in characterizing overall patterns of plant relationships, and emphasizes the uniqueness of plant communities from extrabasinal habitats during the late Paleozoic.     

SPERMOPTERIS,A NEW GENUS OF PTERIDOSPERMS FROM THE UPPER PENNSYLVANIAN SERIES OF KANSAS

Cridland , Arthur A., and John E. Morris . (Kansas U., Lawrence.) Spermopteris, a new genus of pteridosperms from the Upper Pennsylvanian Series of Kansas. Amer. Jour. Bot. 47(10) : 855–859. Illus. 1960.— Spermopteris gen. nov., based upon seed-bearing specimens of the formgenus Taeniopteris, is described from the Lawrence Shale, Pennsylvanian System, of Kansas. The single species S. coriacea (Göppert) comb. nov. is known. Comparison is made with other fertile and supposedly fertile Paleozoic specimens of Taeniopteris and with specimens of T. spatulata from the Rhaetic of Tonkin.