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.     

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.     

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.     

运用数理统计方法对Pecopteris lativenosa Halle 羽叶形态的复原研究

Pecopteris lativenosa Halle是栉羊齿属中羽叶形态变化十分显著的一个种。作者在详细观察的基础上,尝试性地运用定量统计和方差分析方法判定该植物具有代表性的各项特征,提出了普遍存在的5种小羽片类型、6种末次羽片类型和2种末二次羽片类型,以及各类型小羽片、末次羽片、末二次羽片和末三次羽片的空间分布关系,并以此为依据绘制了Pecopteris lativenosa Halle三次羽状复叶复原图。这一复原图能够解释以往发现的Pecopteris lativenosa Halle标本所属类型。     

Osmunda (Osmundaceae) from the Triassic of Antarctica: an example of evolutionary stasis

Compressed specimens of the fern Osmunda are described from the Triassic of the Allan Hills, Antarctica. The specimens consist of a once pinnate, deeply pinnatifid fertile frond as well as several sterile specimens. Six pinnae are present on the partial fertile rachis, with two sterile pinnae above four fertile pinnae. Both sterile and fertile specimens are virtually identical to the modern species Osmunda claytoniana. Entire fronds are fragmentary; the longest is 21 cm in length. Sterile pinnae are alternate and deeply pinnatifid, with slightly toothed pinnules and dichotomous venation. Fertile pinnae are 1-1.3 cm long, once pinnate, and lack vegetative lamina. Sporangia are clustered, each 300-375 um in diameter, and possess a transverse annulus 6-8 cells long; dehiscence is by a vertical slit. Fronds arise from a rhizome 4 cm long by 1 cm wide; two croziers are present on the rhizome. Two frond segments up to 6 cm long and three deeply pinnatifid pinnae are present on the uppermost part of one rachis. Pinnules are ~4 mm long and 2-3 mm wide. The presence of this Osmunda species in the Triassic demonstrates stasis of frond morphology, both fertile and vegetative, for the genus.     

ARCHAEOPTERIS MACILENTA,ANATOMY AND MORPHOLOGY OF ITS FROND

The stem, rachides, and pinnae of Archaeopteris macilenta, formerly considered to be a fern of Devonian age, comprise a branch system in which the ultimate divisions heretofore referred to as pinnules are the leaves. The primary vascular system of the “frond” is a lobed siphonostele from which leaf traces arise in a spiral sequence. The anatomy of the “rachis” and of the “pinnae” is shown to be similar to that of the stem, Callixylon, which bore these “fronds.” Branching, epidermal pattern and stomates are described for the spirally arranged leaves (fertile pinnules). Attachment and dehiscence of sporangia as well as their stomates are reported. Archaeopteris is retained in the Class Progymnospermopsida which includes plants with gymnospermous anatomy and pteridophytic reproduction. It is suggested that Actinopodium, Svalbardia and Siderella are related closely to Archaeopteris and that this group of genera shows evolutionary stages in webbing of leaves and planation of branch systems. The opportunities for ontogenetic studies of the arborescent genus Archaeopteris are pointed out.     

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.     

FOSSIL OPHIOGLOSSALES IN THE PALEOCENE OF WESTERN NORTH AMERICA

Vegetative and fertile frond segments of Botrychium have been recovered from Paleocene deposits of central Alberta, Canada. Specimens are preserved as coalified compressions that yield information about frond structure, sporangia, and spore ultrastructure. These fossils, described as Botrychium wightonii sp. nov., establish a megafossil record for the Ophioglossales, and demonstrate that modern-appearing species of the order were present in western North America by the earliest Tertiary. The largest vegetative fragments are up to 4.6 cm long and tripinnately compound, with opposite to subopposite branching. Ultimate segments are pinnatifid with dentate pinnules and open dichotomous venation. Fertile specimens are also tripinnately compound with a long rachis and subopposite to alternate pinnae. Sporangia are either submarginal and superficial, or marginal, and are all directed toward one surface of the pinnule. They are ovoid to subspheroidal and 0.8-2.0 mm in diameter. Some sporangia are apparently stalked, while others appear to be sessile. This variation results both from the ultimate frond segments being compressed in several different planes, and the fossils being exposed at different levels. Spores macerated from the sporangia are radial and trilete, and range 30–67 μm in diameter. Most are psilate, but some have a densely striate surface.     

The current status of medullosan seed ferns

The compression and permineralized fossil record of presumed medullosan plants is reviewed and evaluated. Recent studies of stelar architecture suggest that stems are monostelic rather than polystelic as commonly believed. Imparipinnate fronds are mostly of the Neuralthopteris/Alethopteris type with Myeloxylon anatomy; the affinity between medullosan stems and neuropterid fronds is uncertain. It is suggested that the reproductive organs of the family should be restricted to those pollen organs believed to have produced Monoletes pollen and to ovules, for the most part, of the Trigonocarpus/Pachytesta type. It is doubtful that reproductive organs replaced lateral pinnules; it is more likely that they terminated pinnae or were borne abaxially along the pinna rachis. In some cases they may have been borne directly on penultimate axes or were, as in Codonotheca and Goldenbergia, borne on fertile fronds or fertile portions of vegetative fronds. Recent research involving pollen organs is emphasized and a phylogeny of Monoletespollen organs is proposed. It is suggested that the fossil record of medullosans is best interpreted from a punctuational perspective.     

STUDIES OF PALEOZOIC FERNS: THE FROND OF ANKYROPTERIS GLABRA

Eggert , Donald A. (Southern Illinois U., Carbondale.) Studies of Palerzoic ferns: The frond of Ankyropteris glabra. Amer. Jour. Bot. 50(4): 379–387. Illus. 1963—The major features of the frond of A. glabra are described on the basis of preserved parts found in Middle Pennsylvanian coal ball material from Illinois. The frond is planated and has well-developed foliar laminae. Primary pinnae arise from the petiole in 2 alternating series, and secondary pinnae arise in a similar fashion from the primary pinnae. Foliar laminae occur on the secondary pinnae and have dichotomous venation. The xylem of the petiole has a diupsilon configuration in the lower part of the axis, while higher in the petiole the xylem forms a strand resembling that of the European species A. westfaliensis. The xylem strands of the primary pinnae arise from the adaxial antennae of the petiolar vascular strand as somewhat C-shapcd bodies and develop antennae and become H-shaped at higher levels. A gap occurs in the antenna of the petiole vascular system above the level of departure of the primary pinna trace. Terete vascular strands occur in the secondary pinna axes which arise from the adaxial antennae of the xylem of the primary pinnae. The foliar laminae are relatively thin, have an irregular outline, and their histology is like that found in many living ferns. The frond of A. glabra illustrates that leaf evolution had progressed in at least one species of the coenopterid family Zygopteridaceae to the extent that an essentially 2-dimensional frond of modern aspect, and with well-developed foliar laminae, was present by Middle Pennsylvanian time.     

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.     

New reconstruction of Weichselia reticulata (Stokes et Webb) Fontaine in Ward emend. Alvin, 1971 based on fertile remains from the middle Albian of Spain

Exceptionally well-preserved specimens from the middle Albian of Spain corresponding to a nearly complete fertile frond and fragments of pinnae containing soral clusters of the tree fern Weichselia reticulata have provided new data about the structure and arrangement of pinnae and their associated fertile reproductive structure. This new material has been compared with the previous studies, and recostructions of this fern and the new data indicate the segregation of vegetative and fertile fronds within the main stipe of this fern.     

Pollen Organ Telangiopsis sp. of Late Devonian Seed Plant and Associated Vegetative Frond

Pollen organ Telangiopsis sp., associated with but not attached to vegetative fronds, has been collected from the Upper Devonian (Famennian) Wutong Formation, Dongzhi County, Anhui Province, China. Fertile axes with terminal pollen organs are dichotomous for 2–4 times and may be proximally attached by fragmentary pinnules. Pollen organs are synangiate and borne on the top of a short stalk. Synangia are radial in symmetry and each consists of 4–8 elongate microsporangia fused at base. Microsporangia have a longitudinal dehiscence line and show a tapered apex. The associated stem is spiny and bears a vegetative frond which bifurcates once at the basalmost part. Frond rachises possess one order of pinna arranged alternately. Pinnules are borne alternately, planate, highly dissected, and equally dichotomous for 2–3 times. Comparisons among Late Devonian seed plants recognize several branching patterns in the fertile fronds/axes bearing terminal pollen organs. Telangiopsis sp. reinforces that the Late Devonian pollen organs are synangiate usually with basally fused microsporangia. It is suggested that the evolutionary divergence of radial and bilateral symmetries of pollen organs may have occurred in the Famennian, when the earliest seed plants evolved planate and sometimes laminate pinnules.     

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.     

Singular taphonomic record of a wildfire event from middle Albian deposits of Escucha Formation in northeastern of Spain

A fossil plant assemblage composed of a great amount of macro, meso and microscopical charcoalifed remains occur in a single layer in Albian deposits of the Escucha Formation in northeastern Spain. This assemblage consists fundamentally of fragments of pinnae and pinnules corresponding to the Matoniaceous ferns Weichselia reticulata and Phlebopteris dunkeri also with some gymnosperm wood remains. The features of both the fossil plants and the deposit itself indicate accurate paleoenvironmental conditions related to the action of wildfires over the vegetation growing in freshwater swamp plains during Albian in Southwestern Eurasia.     

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.     

Fertile and sterile frond segments of the lyginopterid seed fern Feraxotheca

Fertile and sterile frond segments of the lyginopterid seed fern Feraxotheca culcitaus are described. In both the fertile and sterile specimen three orders of frond axes are borne alternately and in the same plane. Antepenultimate and penultimate frond axes are characterized by a C-shaped vascular bundle and numerous canals containing a golden-colored material in the fossilized condition. Synangia with only four sporangia demonstrate radial symmetry; bilateral symmetry is present in larger synangia. Pinnules possess dichotomous venation, with a single vein entering each lobe of the deeply incised lamina. A single canal occurs below each vein, and vein ends have an amplified vascular bundle. Morphologically Feraxotheca compares most closely with the compression genus Crossotheca. The morphology of the vascular bundle in the largest pinna is similar to a Lower Carboniferous specimen of Rhodea.     

THE MORPHOLOGY AND ANATOMY OF CALLIPTERIDIUM SULLIVANTI

Leisman , Gilbert A. (Kansas State Teachers Coll., Emporia.) The morphology and anatomy of Callipteridium Sullivanti. Amer. Jour. Bot. 47(4) : 281—287. Illus. 1960.–Previously undescribed pinnules of Callipteridium Sullivanti (Lesq.) Weiss have been studied in Iowa and Kansas coal balls. Conventional peel techniques revealed the anatomy while deliberately fracturing the coal ball revealed morphological features. Epidermal studies were also conducted after macerating small portions of the coal ball. Evidence indicates that these pinnules were borne on Myeloxylon petioles. Arguments are advanced for removing this species from the genus Alethopteris and returning it to Callipteridium.     

LEAF DEVELOPMENT IN SCHIZAEA AND ACTINOSTACHYS

Observations on the initiation and growth of the fertile digits and of their assemblages in Schizaea and Actinostachys as well as of the entire frond in S. dichotoma are presented. The leaf of S. dichotoma, although dichotomous in appearance at maturity, goes through a series of dichotomous and lateral ramifications during ontogeny. It is compared with the entire frond (“aerial shoot”) of Psilotum nudum. The sporangiate fructifications of Schizaea develop as their mature form suggests, i.e., by acropetal initiation and development of pinnately arranged digits. The counterpart in Actinostachys, irrespective of fertile digit number, develops as only three pinnae, two lateral and one terminal. It cannot be said, therefore, that the condition in Actinostachys, i.e., the digitate arrangement of fertile parts, is fundamentally the condition which exists in Schizaea but with a very short rachis. The fundamental difference is in pinna number and frequency of dichotomies which establish the final digit number.