BOTRYOPTERIS PINNULES WITH ABAXIAL SPORANGIA

Small sporangia borne abaxially on pinnules attached to Botryopteris foliar members are described from coal ball petrifactions of Early Pennsylvanian age. This is the first report of laminar sporangia in this genus. Sporangia are stalked and borne singly near lateral veins on Sphenopteris-like pinnules. Individual sporangia are of the leptosporangiate type, with a lateral annulus and a dehiscence zone of thin-walled cells immediately adjacent to the annulus. Spores are small, trilete, triangular in outline, typically have blunt spines covering the exine, and correspond to the dispersed spore genera Acanthotriletes, Leiotriletes, or Lophotriletes. These sporangia and their spores are unlike previously described globose Botryopteris fructifications from the Middle and Upper Pennsylvanian, but are similar to sporangia produced by modern members of the Osmundaceae.     

AN UNUSUAL BOTRYOPTERID SPORANGIAL AGGREGATION FROM THE MIDDLE PENNSYLVANIAN OF NORTH AMERICA

A new fertile species of Botryopteris (Botryopteridaceae: Filicales) is described from four incomplete Middle Pennsylvanian specimens. Fertile pinnae of B. cratis sp. n. consist of branched frond members bearing numerous globose sporangia. Surrounding the sporangial aggregations are larger sterile frond members (0.5-1.5 mm diam). Fertile pinnae are oval in transverse section and possess an eccentrically developed cortex composed chiefly of fibers. Some frond members show the typical botryopterid xylem configuration with three protoxylem strands. Spherical sporangia are loosely aggregated on the smallest pinnae by short, broad stalks. The annulus is band-like, two cells high, and extends transversely across the lower half of the sporangium for approximately half the circumference. Spores are oval, trilete, verrucate, and covered by a thin separable layer. Sporangium morphology is like that of Botryopteris antiqua, but the spores closely resemble those of B. globosa. The new species is unlike previously described fructifications of Botryopteris in exhibiting a small pinna system which surrounds smaller pinnae bearing sporangia in an aggregation. The new form is considered to be less specialized than previously described globosoid forms because the sporangia are much less crowded. Isolated frond members, believed to belong to the new species, have a large central arm in the pinna xylem trace that resembles the Stephanian taxon B. renaultii. Small stems attached to the adaxial surface of frond members are radial, protostelic, centrarch, and have a three-zoned cortex. The inner cortical zone contains large elongate cells with distinctive layered deposits. Stems are covered with uniseriate multicellular hairs on multicellular bases. Stems compare closely with B. mucilaginosa in histological features.     

The role of Hydropteris pinnata gen. et sp. nov. in reconstructing the cladistics of heterosporous ferns

Large segments of intact plants that represent a heterosporous fern have been discovered within an aquatic plant community from the Late Cretaceous St. Mary River Formation near Cardston in southern Alberta, Canada. Branching rhizomes of Hydropteris pinnata gen. et sp. nov. are 1–2 mm wide. They produce fronds at intervals of 2–12 mm and bear numerous elongated roots. Fronds, up to approximately 6 cm long, are pinnate with subopposite to alternate pinnae that exhibit anastomosing venation. Large, multisoral sporocarps occur at the junctures of the rhizome and frond rachides. Both microsporangiate massulae and megaspore complexes occur within each sporocarp. Megaspore complexes are assignable to the sporae dispersae genus Parazolla Hall. Microspores are trilete, smooth-walled, and are embedded in episporal material of the massulae. A numerical cladistic analysis indicates that the heterosporous aquatic ferns are monophyletic, and not as closely related to either schizaeaceous or hymenophyllaceous ferns as they are to some other filicaleans. Systematic revisions are proposed to reflect newly recognized cladistic relationships within the heterosporous clade, and character originations in the evolution of heterosporous aquatic ferns are evaluated. Hydropteridaceae fam. nov. is proposed, and included with Salviniaceae and Azollaceae in the Hydropteridineae subord. nov., and the Hydropteridales Willdenow.     

RHABDOXYLON AMERICANUM SP. N., A STRUCTURALLY SIMPLE FERN-LIKE PLANT FROM THE UPPER PENNSYLVANIAN OF ILLINOIS

A new species of the genus Rhabdoxylon Holden (1960), an anatomically simple plant of presumed fern or fern-like affinities, is described from a coal ball petrifaction found in the Upper Pennsylvanian of southern Illinois. The new species, R. americanum, is based upon five specimens consisting of stems bearing spirally arranged leaves and numerous randomly distributed adventitious roots. The haplostelic stems branch by equal dichotomies and bear foliar traces which arise as unequal dichotomies of the stele. Leaf traces possess a circular outline in cross section and one adaxial protoxylem strand. The characteristics of exclusively primary tissues, diarch adventitious roots, centrarch haplosteles with simple scalariform pitting, and the nature and arrangement of the leaf traces, suggest that Rhabdoxylon represents a fern or fern-like plant rather than a representative of the Rhyniophytina or Trimerophytina. At present it is not possible to determine whether the simple structure of Rhabdoxylon has come about through phyletic reduction or represents a primitively simple condition.     

A PENNSYLVANIAN CALAMITEAN CONE WITH ELATERITES TRIFERENS SPORES

In 1943 L. R. Wilson described some scattered elater-bearing spores found in a Middle Pennsylvanian coal ball from What Cheer, Iowa, as Elaterites triferens. The spores, averaging 60 μ in diameter, with a trilete scar on the proximal surface and three conspicuous elaters attached to their distal surface, have now been found in a fragment of a cone from the same locality. The cone is similar to Calamostachys in having whorls of sporangiophores, each with four adaxial sporangia, and some secondary xylem at the nodes, but it possibly differs in lacking sterile bracts.     

CALAMOCARPON INSIGNIS,A NEW GENUS OF HETEROSPOROUS,PETRIFIED CALAMITEAN CONES FROM THE AMERICAN CARBONIFEROUS

Baxter , R. W. (U. Kansas, Lawrence.) Calamocarpon insignis , a new genus of heterosporous, petrified calamitean cones from the American Carboniferous. Amer. Jour. Bot. 50(5): 469–476. Illus. 1963.—Calamocarpon insignis is described as a new genus of calamitean cone based on numerous fragments of microsporangiate and megasporangiate cones found in coal balls of middle Pennsylvanian age from the Cherokee Group of Kansas and the Des Moines Series of Iowa. The cones are similar to Calamostachys in general construction, having a hollow pith, prominent protoxylem canals, and alternating whorls of sterile bracts and sporangiophores. They differ in that the microsporangiate cones bear microsporangia each containing several hundred microspores which may occur as tetrads or single spores, while the megasporangiate cones bear large rectangular megasporangia each containing a single functional megaspore surrounded by sterile nutritive tissue. The microspores average 30–40μ in diameter compared to a maximum measurement of 2.7 × 0.7 mm for the single rectangular megaspore. The female gametophyte was produced within the megaspore which was held within the megasporangium during the entire period of development. The megasporangia were deciduous so that the entire structure was shed as a unit.     

VEGETATIVE AND FERTILE STRUCTURES OF CYATHOTHECA VENTILARIA FROM THE UPPER PENNSYLVANIAN OF THE APPALACHIAN BASIN

Permineralized stems, leaves and a fertile structure assignable to Cyathotheca Taylor are described from the Late Pennsylvanian Duquesne Coal of eastern Ohio. The new material, C. ventilaria sp. nov., provides the first evidence of vegetative structures for the genus. Vegetative parts are referred to the fertile fragments based upon distinctive vascular morphology, common histological features, and close association. Stems are up to 5.0 mm in diam and have an apparently endarch dictyostele with scalariform metaxylem tracheids. Secondary xylem consists of tracheids with bordered pitting and uniseriate, parenchymatous rays 1–9 cells high. Leaves are arranged in a 3/8 pattern. They are small, pinnately-lobed, and vascularized by an U-shaped bundle. Distal to divergence, one of the apparent leaf traces becomes radial to resemble the base of a fertile structure. This implies a mode of attachment for the fructification and suggests evidence to interpret its homologies. The fertile specimen consists of laminae that diverge from a short stalk, and bifurcate distally. Laminae bear sporangia 0.6–0.7 mm in diam attached adaxially by vascularized pedicels. Spores average 36 μm in diam and are of the Kewaneesporites type. The combination of features now known for Cyathotheca exclude it from assignment to a currently-recognized major group of vascular plants, thus emphasizing that the Pennsylvanian coal swamp flora included a greater diversity than commonly is interpreted.     

FERAXOTHECA GEN. N., A LYGINOPTERID POLLEN ORGAN FROM THE PENNSYLVANIAN OF NORTH AMERICA

The pollen organ Feraxotheca gen. n. is described from Pennsylvanian age coal balls from the Lewis Creek, Kentucky, locality. The fructifications consist of bilaterally symmetrical synangia composed of a basal pad supporting elongate sporangia that are laterally appressed for the entire length of the sporangial cavities. Sporangial tips extend over the center of the synangium and delimit a small open area, while the bases arise from a parenchymatous cushion that is bounded by short tracheid-like cells. Each synangium is borne on the surface of an expanded pinna tip and is surrounded by a small amount of laminar tissue that envelopes the base of the synangium. Ultimate pinnae are rectangular in transverse section, possess an elliptical vascular bundle surrounded by canals containing a yellow froth-like substance, and have a cortex of elongate cells that radiate from the center of the axis. Sporangia contain small (40–64 μm), radial, trilete spores ornamented by regularly spaced coni or blunt tipped grana. Feraxotheca is compared with the compression genus Crossotheca and some new ideas are advanced concerning the morphology of this compression genus. The obvious differences between Feraxotheca and other lyginopterid pollen organs strongly suggests that the Lyginopteridaceae, as it is currently interpreted, is an unnatural family.     

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.     

Paratingia wudensis sp. nov., a whole noeggerathialean plant preserved in an earliest Permian air fall tuff in Inner Mongolia, China

Noeggerathiales are a little known group of Carboniferous and Permian plants of uncertain systematic position that have been variously considered to be ferns, sphenopsids, progymnosperms, or a separate group. These heterosporous plants carry adaxial sporangia on leaf-like or disk-shaped sporophylls that form cones. Leaves are pinnate with a rather stiff appearance, and pinnules can be attached in either two or four rows. In the present report, we present the top of a noeggerathialean plant with leaves and strobili attached, Paratingia wudensis Wang, Pfefferkorn et Bek sp. nov., from an earliest Permian volcanic ash fall tuff in Inner Mongolia. The excellent preservation allows the reconstruction of the whole plant, the complex three-dimensional leaves with anisophyllous pinnules, the heterosporous strobili, and the spores in situ. The homology of leaves and strobili can be elucidated and contributes to an understanding of the debated taxonomic position of Noeggerathiales. The "anisophyllous" leaves carry pinnules arranged in four rows. The strobili are bisporangiate and have disk-shaped sporophylls, each with one ring of 10-14 adaxial sporangia around the strobilus axis. Megaspores have an equatorial bulge. This new species expands the known diversity of Noeggerathiales. It grew in a peat-forming forest, thus changing earlier interpretations of the growth of noeggerathialean plants with anisophyllous pinnules.     

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.     

STUDIES OF PALEOZOIC MARATTIALEANS: THE MORPHOLOGY AND PROBABLE AFFINITIES OF TELANGIUM PYGMAEUM GRAHAM

Telangium pygmaeum Graham is known from Upper Pennsylvanian coal balls from the Calhoun coal mine (Illinois). The species was described as possessing radial synangia consisting of 3-5 sporangia fused laterally for about f13 their length. Synangia were believed to be sessile and borne terminally or laterally on a branching rachis without lamina. Examination of new coal ball material of the same age indicates that the synangia are borne abaxially on the pinnules of a compound frond with the anatomy of a Psaronius leaf (Marattiales). Synangia are sessile and borne in two rows, one on either side of the pinnule midrib, under the unbranched lateral veins. Synangia are radial, 0.6 mm in diam, and consist of a ring of thin-walled sporangia fused to near their apices prior to dehiscence, but separating on dehiscence to release spores along their inner midline. Spores are spherical, trilete, 30-48 μm in diam, with a granulate ornamentation. The new genus Araiangium is proposed for this material based on the organization of the sessile thin-walled synangia. Araiangium is compared with other marattialean genera with sessile synangia (Acaulangium, Acitheca), and with the pedicellate synangia of various species of Scolecopteris. Criteria used in the delimitation of genera in Paleozoic anatomically preserved marattialean fertile foliage are discussed.     

SULLITHECA DACTYLIFERA GEN. ET SP. N.: A NEW MEDULLOSAN POLLEN ORGAN AND ITS EVOLUTIONARY SIGNIFICANCE

The pteridosperm (Medullosaceae) pollen organ Sullitheca dactylifera gen. et sp. n. is described from middle Pennsylvanian coal balls. The proximally fused units of the obpyriform compound synangium separate and extend distally as finger-like projections. Each projection contains 4–6 vertically oriented cylindrical sporangia arranged in pairs along the radius of the unit; each unit extends from the outer cover wall toward the center. The distal portion of the compound synangium is hollow as a result of the lateral separation of the centripetally and distally directed synangial units. About 40 tubular sporangia are present in all and dehiscence occurs along a lateral slit in each sporangium. Vascular strands are disposed around the periphery of the organ in addition to a single strand paralleling each sporangium. Two- or three-cell trichomes and stomata are present on the organ surface. Pollen of the Monoletes type is present. A paired row of sporangia in Sullitheca composing a synangial unit is considered the homologue of a paired row of sporangia in the more compact and highly evolved genus, Dolerotheca.     

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.     

CALLANDRIUM CALLISTOPHYTOIDES,GEN. ET SP. NOV., THE PROBABLE POLLEN-BEARING ORGAN OF THE SEED FERN,CALLISTOPHYTON

A new genus of pteridosperm pollen organ is described from Pennsylvanian age coal balls of Illinois. Individual sporangia are grouped into radial synangia which are borne in opposite pairs on the abaxial surface of slightly modified pinnules. Sporangia contain monosaccate pollen with a distinct sulcus, referable to the pollen genus Vesicaspora. Pinnules are borne on a regularly pinnate frond which is circinately coiled when young. At least a large portion of the frond is fertile and possesses anatomical features similar to those of Callistophyton. Phyletic relationships with other pteridosperm pollen organs are discussed.     

POROSTROBUS NATHORSTII SP. NOV.: A NEW SPECIES OF LYCOPSID CONE FROM THE EARLY PENNSYLVANIAN OF ILLINOIS

Thirty-one specimens of a small megasporangiate lycopsid cone referable to the genus Porostrobus Nathorst and abundant associated dispersed megaspores have been collected from Early Pennsylvanian strata in the Allied Stone Company quarry, Milan, Illinois. Based on other elements in the flora, the deposit is considered to be part of the Morrowan Caseyville Formation and probably of Namurian age. This is the first reported occurrence of Porostrobus in North America and the cones are recognized as a new species, P. nathorstii. The environment of deposition indicates that the cones may have been transported from the parent plant prior to preservation. Cones are preserved as coalified compressions measuring 15–36 mm long by 2.5–7 mm wide, and are characterized by an apical tuft of leaves up to 20 mm long. Sporophylls are spirally arranged on a narrow cone axis, lack a heel or keel, and have a long distal lamina. Sporangia contain a single functional megaspore tetrad. Mature megaspores are 750–1, 150 μm in diameter, have prominent trilete sutures raised to form a gula, and have numerous branched hairs confined to an equatorial band. Megaspores correspond to the dispersed form Setosisporites praetextus (Zerndt) Potonie and Kremp. Porostrobus nathorstii is the only species of the genus described to date that is monosporangiate.     

THE SPORANGIUM OF HORNEOPHYTON LIGNIERI (RHYNIOPHYTINA)

Numerous sporangia of Horneophyton lignieri from the Rhynie Chert locality in Scotland have been studied. The sporangia are branched, with two to four columellate lobes of varying length, and a continuous sporogenous zone or cavity occurs among the lobes. Unbranched sporangia, generally thought to be the typical form for the plant have not been found, and their presence is not established. Although not definitely proven, evidence suggests that the sporangia opened by means of a small apical pore or stoma. An area of thick-walled cells at the apex of each sporangial lobe probably played some role in this opening. Radial, trilete, azonate spores ranging from 39–49 μm in diam, with curvaturae perfectae are produced most commonly in tetrahedral tetrads and occasionally in isobilateral tetrads. Matters of spore preservation and possible ornamentation are discussed. The branched sporangia of this genus are unique among bryophytes and vascular plants and provide some evidence that certain synangia may have arisen from a single sporangium rather than from multiple sporangia borne singly at the tips of ultimate branches.     

A NEW SPECIES OF ARTHROXYLON (CALAMITACEAE) FROM THE PENNSYLVANIAN OF KANSAS

Cridland , Arthur A. (Kansas U., Lawrence.) A new species of Arthroxylon (Calamitaceae) from the Pennsylvanian of Kansas. Amer. Jour. Bot. 46(10): 709–712. Illus. 1959.—Arthroxylon resinaceum sp. nov. is described from 3 specimens found in a coal-ball collected at West Mineral, Kansas. The tracheids have 1 or 2 rows of pits on the radial walls and the pith cells adjacent to the protoxylem canals are filled with brown contents. Fungus spores are present in the tissues of 1 specimen.     

STUDIES OF PALEOZOIC MARATTIALEANS: AN EARLY PENNSYLVANIAN SPECIES OF THE FERTILE FERN SCOLECOPTERIS

Remains of the fossil Marattiales are very rare in Lower Pennsylvanian sediments. The present report describes a new species of the fertile fern foliage Scolecopteris from the Lewis Creek, Kentucky locality (Lower or lower Middle Pennsylvanian). Scolecopteris conicaulis n. sp. has radial synangia composed of a ring of 4–7 elongate, exannulate sporangia. Most features of the synangia of S. conicaulis were previously hypothesized to be primitive in Scolecopteris based on geologically younger species. Supposed primitive characters include the large synangium pedicel with fiber core, an outer-facing sporangial wall lacking differentiation or zonation, and large spores. The anatomy of the sporangium walls, pinnule morphology, and general spore type support an association with the Minor group of Scolecopteris. The new species is similar in several important features to Scolecopteris (Cyathotrachus) altus, the only other anatomically preserved fertile marattialean known from this early time, and indicates a considerably earlier origin for fertile foliage of this type.     

PALEOZOIC SEED FERNS: HETERANGIUM KENTUCKYENSIS SP. NOV., FROM THE UPPER CARBONIFEROUS OF NORTH AMERICA

Heterangium kentuckeyensis sp. nov. is described from Lower/Middle Pennsylvanian sediments of eastern Kentucky, based on permineralized stems, petioles, frond members, laminar foliage, and roots, including several organs in attachment. Stems 2.2 to 5.8 mm in diameter are known in several developmental stages. The considerable variability in stelar and cortical histology within this one species emphasizes the need to reassess variability within previously described taxa. Fronds of H. kentuckyensis are at least twice pinnate and bear primary pinnae alternately at approximately right angles. Laminar pinnules have dichotomous venation, are at least 2-lobed, and comparable to foliage of the Sphenopteris-type. Stomata possess 6–7 subsidiary cells with abaxial papillae. The characters used to distinguish the subgenera of Heterangium are evaluated and found to be unreliable. Moreover, suggested phylogenetic schemes both within Heterangium and between this taxon and other lyginopterid pteridosperms based on these features are inconsistent with stratigraphic data. Until reproductive features are known, the classification of Heterangium species is best based on characters of the vegetative sporophyte, including stelar organization (particularly protoxylem architecture), cortical histology, and frond morphology.