Fungal sporocarps from the Carboniferous: An unusual specimen of Traquairia

Small spherical structures commonly termed sporocarps are widespread elements in Carboniferous permineralized peat. While sporocarps today are generally interpreted as fungal, their precise systematic affinities largely remain unknown. A specimen of the sporocarp-type Traquairia from the Lower Coal Measures (Lower Pennsylvanian) of Great Britain demonstrates a heretofore unknown feature in the form of a preformed aperture from which emerges a fascicle of radially oriented structures that are constricted sub-distally; transverse septa are present in the constricted areas of some of the structures. The outgrowths are morphologically similar to conidiophores bearing terminal conidia of certain extant fungi in the order Entomophthorales, and thus might suggest affinities of Traquairia with the Zygomycota. Alternatively, the outgrowths might represent sporulating hyphae of a parasitic or saprotrophic blastocladalean fungus or a peronosporomycete that used Traquairia as a nutrient source.     

STUDIES OF PALEOZOIC FUNGI. III. FUNGAL PARASITISM IN A PENNSYLVANIAN GYMNOSPERM

Evidence of fungal parasitism is found in the Pennsylvanian gymnospermous cone, Lasiostrobus polysacci Taylor. Indication of fungal activity is found in the outer cortical region of the axis of the cone and in the fleshy microsporophylls. Specimens exhibit severe tissue disruption, thick-walled, branched, septate hyphae, and possible reproductive structures. Parenchymatous cortical cells may also contain rounded bodies which are continuous with the cell wall. Similar structures are formed in many extant taxa in response to fungal invasion, and are termed wall appositions or callosities. Although their role in extant plants is disputed, they are clearly the product of a living host cell. Such spherical bodies, however, are not restricted to the cell periphery but in some cases occlude the cell lumen. In appearance they resemble resinous remains similar to those found in other coal ball plants. The blockage of entire cells or groups of cells may have served to retard hyphal growth or isolate infected cells. The occurrence of such structures in a Carboniferous plant provides the best evidence to date of parasitism during the Paleozoic.     

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.     

FUNGAL REMAINS IN THE DEVONIAN TRIMEROPHYTE PSILOPHYTON DAWSONII

Fungal reproductive structures were found within aerial axes of the Late Devonian trimerophyte, Psilophyton dawsonii Banks, Leclercq and Hueber from Quebec, Canada. Specimens are spherical to oval, up to 175 μm in maximum diam, and exhibit two distinct wall layers. They are restricted to the inner cortical region of infected axes. The fungal bodies resemble species of Palaeomyces from the well-known Rhynie Chert flora, and are also similar to the chlamydospores commonly found in Pennsylvanian coal ball plants. Among extant organisms they are most similar to oomycetous oogonia and endogonaceous chlamydospores.     

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.     

STUDIES OF CARBONIFEROUS FUNGI. II. THE STRUCTURE AND ORGANIZATION OF MYCOCARPON,SPOROCARPON, DUBIOCARPON,AND COLEOCARPON (ASCOMYCOTINA)

The problematic organism, Sporocarpon, was among the first fossil fungi to be described. The genus was actually a heterogeneous assemblage of fungal forms, and was eventually divided into several taxa including Sporocarpon, Dubiocarpon, and Mycocarpon. In the present paper these organisms, as well as a similar, previously undescribed genus, Coleocarpon; are described from Lower, Middle, and Upper Pennsylvanian strata in North America, and from Upper Carboniferous sediments in England. Although initially considered to be radiolarian-like protozoa, these structures show far more similarity to ascomycetous cleistothecia. All forms are ornamented, spherical bodies with a complex wall which encloses scattered asci and ascospores. Taxa are distinguished primarily by differences in the organization of the cleistothecial wall. Structurally similar organisms are found today among the Eurotiales.     

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.     

CUTICLES OF TWO SPECIES OF ALETHOPTERIS

Specimens of Alethopteris sullivanti (Lesquereux) Schimper and Alethopteris lesqueieuxi Wagner were found in Middle Pennsylvanian coal balls from the Lovilia, Iowa and West Mineral, Kansas localities. Light microscopy and scanning electron microscopy (SEM) were used to study macerated cuticles of these two foliage types of the pteridosperm, Medullosa. The lower epidermises of both species of Alethopteris possess sunken stomata, papillae, and evidence of multicellular hairs. The upper epidermises are devoid of these features. Critical point drying and subsequent observation with SEM were used to document the structure of the multicellular hairs of A. lesquereuxi. Replicas of the lower epidermal features of an impression-compression specimen of A. sullivanti were observed with SEM and compared with the petrified specimens.     

STUDIES OF PALEOZOIC SEED FERNS: ANATOMY AND MORPHOLOGY OF MICROSPERMOPTERIS APHYLLUM

The discovery of numerous specimens of the monostelic pteridosperm genus Microspermopteris in Pennsylvanian coal ball petrifactions from the Lewis Creek and What Cheer localities provides additional information about the anatomical and morphological variability within the genus. Specimens are now known up to 1.1 cm in diam that bear epidermal appendages in the form of variously-shaped trichomes. The external surface of the stem is further ornamented by longitudinal flaps of cortical tissue. Petioles exhibiting a single C-shaped vascular strand with abaxial protoxylem are produced in a 2/5 phyllotaxy. Large petiole bases that clasp the stem produce primary pinnae alternately. The presence of axillary branching appears similar to that reported in Callistopliyton and Lyginopteris. Triarch to polyarch adventitious roots, some with secondary tissues, are produced at both nodal and internodal regions. Of the currently recognized monostelic seed fern genera, Microspermopteris is most similar to Heterangium. Information is presented that supports current ideas regarding the evolution of the gymnospermic eustele from protostelic Devonian ancestors.     

THE FROND OF HETERANGIUM

Frond members belonging to the monostelic seed fern genus Heterangium have been found in Pennsylvanian age coal balls collected in the Illinois Basin. Petioles bear small pinnae below a dichotomy which produced a bipartite frond. Pinnules of the Sphenopteris obtusiloba type are borne on secondary pinnae. The anatomy of each pinna order and the pinnules is described.     

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.     

BOTRYOPTERIS FORENSIS (BOTRYOPTERIDACEAE), A TRUNK EPIPHYTE OF THE TREE FERN PSARONIUS

Basal parts of Botryopteris forensis have been discovered rooted within the mantle of the tree fern Psaronius. Specimens occur in Upper Pennsylvanian coal balls from near Steubenville, Ohio, USA. The Botryopteris stems branch profusely, and these shoots are intertwined with the Psaronius roots near the surface of the mantle. They also produce adventitious roots that extend among the Psaronius roots. This material demonstrates that B. forensis was a trunk epiphyte, rather than a rhizomatous terrestrial fern. The B. forensis plant is interpreted to have branched continuously, to ramify, and to maintain itself at the periphery of the growing mantle of Psaronius roots. A new reconstruction of B. forensis is offered showing the large, globose fructifications hanging pendulously from horizontal fronds on emergent shoots. Epiphytes and lianas are common on the trunks of Psaronius, indicating that some Marattiales did not produce leaf skirts.     

PROBABLE ATTACHMENT OF THE DOLEROTHECA CAMPANULUM TO A MYELOXYLON-ALETHOPTERIS TYPE FROND

The discovery of pedunculate specimens of pollen organ Dolerotheca in close association with Myeloxylon-type pinnae and Alethopteris-type pinnules provides evidence for the reconstruction of a petrified medullosan frond and the attachment of the microsporangiate fructification. Specimens of Dolerotheca villosa and D. formosa from Middle and Upper Pennsylvanian age strata of North America are borne on slender peduncles with anatomical features and vascularization identical to those of some level in a Myeloxylon-Alethopteris-type frond. Reconstruction of the frond reveals a large, repeatedly dichotomising organ that bears penultimate pinnae and ultimate pinnules in a pinnate arrangement. Examination of pinnules on the surface of coal ball material indicates that they conform to the compression genus Alethopteris. Identical numbers and arrangement of vascular bundles together with identical anatomical features and multicellular hairs indicate that the Dolerotheca campanulum is borne in the position of a penultimate pinna. A reconstruction of the frond with a proposed polled organ attachment is included.     

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.     

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 ANATOMY AND THREE-DIMENSIONAL MORPHOLOGY OF ANNULARIA HOSKINSII SP. N.

Foliage attached to calamitean shoots is described from coal ball petrifactions of Middle and Late Pennsylvanian age. Leaves correspond to the compression-impression genus Annularia and thus represent the first attached members of this genus to be recognized as petrifactions. Individual leaves contain a single unbranched vascular bundle flanked by wide lateral laminar areas which occupy more than half the leaf cross sectional area. Stomata are confined to broad bands within concave furrows which parallel the vascular bundle on the abaxial leaf surface. Epidermal cells within these furrows are in rows aligned obliquely to the leaf axis, and the rows angle outward at a slight angle towards the leaf margin. Convolutions of the leaf-bearing axes result from nodal diaphragms which are oblique to the shoot axis. Whorled leaves apparently radiate outward in the plane of each obliquely positioned nodal diaphragm. This petrified material helps explain the apparent flattening of entire nodal diaphragms and leaf whorls within the same plane as seen in compression specimens. Annularia hoskinsii sp. n. is proposed, and the systematic position of structurally preserved Annularia foliage relative to the genus Dicalamophyllum is discussed.     

POLLEN ORGANS OF THE PENNSYLVANIAN CALLISTOPHYTACEAE (PTERIDOSPERMOPSIDA)

Numerous specimens of the pteridospermous pollen organs Idanothekion glandulosum and Callandrium callistophytoides are present in Middle and Upper Pennsylvanian petrifaction material, respectively. Several specimens of Idanothekion, previously known only in an isolated condition, are attached to foliage assignable to the monostelic seed fern Callistophyton. Proof of the affinities of Idanothekion allows this genus to be elevated from a form genus to an organ genus in the Callistophytaceae. The structure of the foliage and the mode of attachment of the Idanothekion specimens are like those previously described for Callandrium. Structural features of Idanothekion and Callandrium are compared, and the question of their identity is discussed. At present one basic type of pollen organ, borne on the abaxial surface of foliar pinnules, is known for members of the Callistophytaceae.     

ONTOGENY OF THE PALEOZOIC OVULE,CALLOSPERMARION PUSILLUM

The ontogeny of the upper Pennsylvanian age gymnospermous ovule, Callospermarion pusillum, is described from petrifaction specimens collected at the Berryville locality in Illinois. Ovules exhibit a wide range of dimensional and structural features that indicate an extensive developmental sequence. Specimens range from ovules with indistinct zonation of the thin-walled integument to those with thick-walled cells of the sclerotesta. The apex of the fleshy nucellus in some specimens is preserved as a cellular mound, while in others a well-formed cellular pollen chamber is present; still other ovules are characterized by a papery-thin nucellus and pollen chamber wall. The megagametophyte of most specimens is represented by a hollow megaspore membrane that may be restricted to the base of the nucellus, or occupy the entire seed cavity. In a few specimens cellular gametophytes are preserved, and in one ovule archegonia with supposed eggs are also present. Variability in each of the features is compared with ontogenetic changes in comparable structures of living gymnospermous ovules and is correlated with ovule size. A developmental sequence for the fossil ovules is proposed.