ACAULANGIUM GEN. N., A FERTILE MARATTIALEAN FROM THE UPPER PENNSYLVANIAN OF ILLINOIS

Material described by Graham as Cyathotrachus bulbaceus is believed to represent a new genus that is a common constituent of Upper Pennsylvanian coal balls. The sessile synangia of Acaulangium gen. n. are borne in a row on either side of the pinnule midrib and are composed of four to six short, tapering, laterally appressed sporangia. The sporangia have extended tips which curve over the inside of the synangium distally and delimit a small open area inside the synangium. The outer facing walls of the sporangia are two to three cells thick throughout while the inner facing walls are uniseriate. During dehiscence the sporangia separate laterally and spore release results from the rupture of a row of elongate cells along the inner sporangium midline. Among species of Scolecopteris the new genus resembles S. illinoensis and S. minor var. parvifolia but differs in its sessile synangial attachment. The additional parenchyma present between sporangial cavities in the synangia of Acaulangium, and the tendency toward bilateral symmetry suggests an early stage in the evolution of a bivalve synangium such as is present in Marattia.     

IDANOTHEKION GEN. N., A SYNANGIATE POLLEN ORGAN WITH SACCATE POLLEN FROM THE MIDDLE PENNSYLVANIAN OF ILLINOIS

Idanothekion glandulosum gen. et sp. n. is a synangiate pollen organ represented by approximately 30 specimens contained in coal balls from the middle Pennsylvanian of Illinois. Each synangium is composed of seven to nine elongate sporangia that are fused laterally for approximately four-fifths of their length, and are radially arranged about, and fused to, a short central column; the central column is restricted to the proximal one-third of the synangium. Distal to the column the sporangia surround a hollow central area. Dehiscence occurred by means of a longitudinal slit along the mid-line of the inner face of each sporangium. The outer walls of the sporangia have a complex histology involving an external epidermis, a middle presumably glandular layer containing scattered enlarged cells, and an inner layer made up of thin-walled parenchyma. Vascular tissue is present in the central column and outer walls of the sporangia. Each sporangium has a prominent, attenuate, multicellular tip. Large numbers of saccate pollen grains similar to those found in numerous fossil and extant coniferophytes as well as some Mesozoic pter-idosperms were produced in each sporangium. Idanolhekion resembles some synangia assignable to Paleozoic members of the Marattiales; however, the new genus compares most closely with pollen organs believed to have been produced by members of the Pteridospermales. It seems most likely that Idanothekion represents the pollen organ of some member of the Lyginopteridaceae that produced pollen of a type which up to now has not been known from Paleozoic seed ferns.     

Melissiotheca: a new pteridosperm pollen organ from the Lower Carboniferous of Scotland

Melissiotheca is based on fusainized pollen organs of pteridospermous affinities which occur abundantly in an Upper Visean limestone at Kingswood, near Pettycur (Scotland). The new species is a pedicellate synangium composed of 50–150 sporangia each of which is embedded at its base in a parenchymatous cushion divided into lobes. The sporangia are fused along their proximal half but are free distally. Dehiscence is longitudinal. Each sporangium is supplied at its base by a single vascular strand. Prepollen is small, spherical and trilete with a rugulate exine. In ultrastructure, the nexine appears homogeneous; the sexine shows internal sculpture of granae and rod elements. Melissiotheca has not been assigned to any family, but it shows many affinities with pollen organs attributed to the Lyginopteridaceae.     

Aulacotheca collicola n.sp. (medullosaceae) from the early Pennsylvanian of the Illinois basin

Compressed synangia referable to the medullosan pollen organ genus Aulacotheca Halle have been discovered in Early Pennsylvanian sediments from Rock Island County, Illinois, U.S.A. Specimens were recovered from localized shales containing floral elements suggesting an upland environment. Based on synangial size, morphology, sporangial number, and prepollen type, two distinct forms, A. collicola n.sp. and Aulacotheca sp., are recognized. Synangia of A. collicola are 11–17 × 2.5 – 4.0 mm and have four to six sporangia. Free sporangial tips are acutely pointed and dehiscence is through a longitudinal slit to the inside of the synangium. Prepollen averages 181 × 129 μm and has a monolete suture with median deflection. Sexine on proximal and distal surfaces shows narrow, closely spaced, anastomosing inpockets; distal and lateral walls are separated by a deep, encircling fold. Synangia of Aulacotheca sp. are 21–33 × 3.8 – 5.5 mm and have 4–6 sporangia. Distally, free sporangial tips have a narrow extension, giving a mucronate appearance. Features of these species suggest that greater morphological diversity in synangial and pollen structure occurs in Aulacotheca and the medullosan pteridosperms than previously recognized.     

Melissiotheca: a new pteridosperm pollen organ from the Lower Carboniferous of Scotland

Melissiotheca is based on fusainized pollen organs of pteridospermous affinities which occur abundantly in an Upper Visean limestone at Kingswood, near Pettycur (Scotland). The new species is a pedicellate synangium composed of 50–150 sporangia each of which is embedded at its base in a parenchymatous cushion divided into lobes. The sporangia are fused along their proximal half but are free distally. Dehiscence is longitudinal. Each sporangium is supplied at its base by a single vascular strand. Prepollen is small, spherical and trilete with a rugulate exine. In ultrastructure, the nexine appears homogeneous; the sexine shows internal sculpture of granae and rod elements. Melissiotheca has not been assigned to any family, but it shows many affinities with pollen organs attributed to the Lyginopteridaceae.     

DIMENSIONAL CORRELATIONS IN DEVELOPING SELAGINELLA SPORANGIA

With length of sporangia as a developmental index, the growth relationships of sporangia during differentiation were studied in strobili of Selaginella bigelovii. The strobili usually contain two rows of megasporangia and two rows of microsporangia with a mega- opposite a microsporangium at each node. Prior to the sporocyte stage a sporangium in a megasporangiate row is larger and elongates more rapidly than a sporangium opposite it at the same node in a microsporangiate row. The number of sporogenous cells is similar in sporangia of the same length from both rows until cell multiplication ceases in sporangia of the megasporangiate row, while it continues in sporangia of the same size in the microsporangiate row. The observed growth differences between sporangia of the micro- and megasporangiate rows are interpreted as events in the differentiation of two sporangial types.     

Anatomy of a New Species of Scolecopteris (Marattiales) from Early Stage of Early Permian in China

Scolecopteris Zenker, a kind of anatomically-preserved fertile foliage of Late Paleozoic Marattiales, has been well studied in Euramerican Flora. It is composed of 28 species which can be divided into four forms (groups) mainly based on modified or umodified pinnules, the variation of the outer facing sporangial walls, and with or without a prominent central parenchyma area. In contrast, Scolecopteris Zenker in Cathaysian Flora has been poorly studied so far, and only one species S. sinensis Zhao, was reported in 1991 which was considered as a member of Minor Group. The paper reports a second species of Scolecopteris, i.e.S, shanxiensis sp. nov., which differs from the above four groups in that its outer facing wall of the sporangia is thick at the base and top (2 ~ 3 layers of cells), and a little thinner ( 1~2 layers of cells) at the midlevel of the synangia. So a new group, Shanxiensis group, is set for the new species. The other characteristics of Shanxiensis group is comparable with Minor group. The new species comes from the coal balls in Coal Seam No. 7 in the upper part of Taiyuan Formation (early Early Permian) from Taiyuan, Shanxi, China. The identification of Scolecopteris shanxiensis sp. nov. :The fertile pinnule probably peeopterids, 5.5 ~ 6.0 mm in length and 2.0 ~ 2.2 mm in width. The lateral extensions of the lamina of the pinnule bend abaxially and above the synangia. The synangia arrange along the sides of the midrib of the pinnnle and there are about 10 synangia in each row. The synangium is elliptical in longitudinal section and radial in cross section, 0.7 ~ 0.8 mm in height and 0.6 ~ 0.7 mm in diameter. Each synangium has 5-7 (mostly 6) fusiform sporangia fused at the base and attached to the top of the synangial pedicel. The outer facing wall of the sporangia consists of 2~3 layers of cells at the base and becomes thinner at the midlevel (1~2 layers of cells), and at the top of the synangia the wall become thicker again. The cells of the outer facing wall of the sporangia are elongate in the longitudinal sections. The lateral and inner facing walls of the sporangia are one cell thick. The synangial pedicel is small. Spores in situ are small, generally 11~14 µm in diameter, spherical or rounded-triangular, trilete and smooth-walled.     

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.     

STEWARTIOTHECA GEN. N. AND THE NATURE AND ORIGIN OF COMPLEX PERMINERALIZED MEDULLOSAN POLLEN ORGANS

Stewartiotheca gen. n. is a bell-shaped, unisynangiate pollen organ with eccentric radial symmetry and a single series of about 80 pollen sacs. Infoldings that vary in depth occur circumferentially and extend from the periphery to a point off center. This position also marks the location of a sclerenchyma column (proximally) and a sclerenchyma-lined, conical hollow (distally) that opens onto the distal face of the organ. Plates of ground parenchyma extend inward from the outer covering of the organ at locations of infoldings, while similar plates with sclerenchyma strands occur between these locations. Pre-pollen of Monoletes type was released through distal longitudinal slitlike openings of the pollen sac faces toward the sclerenchymatous ground tissue plates. Vascular bundles entering the organ undergo repeated dichotomies, and lead to numerous bundles both in the cover (one per sac for those sacs that abut directly on cover tissue) and internally (one per pair of pollen sacs that lie opposite one another across the location of an infolding). The most complex permineralized medullosan pollen organs Sullitheca, Stewartiotheca, and Dolerotheca are considered to have evolved from a similar type of cup-shaped organ with a single ring of pollen sacs, broadly open distally, and with a central hollow. Circumferential infoldings of one organ of this type were involved in the origin of both Stewartiotheca and Sullitheca, while four similar organs, each showing infoldings non-circumferentially, fused to produce the Dolerotheca type organ (exemplified by D. formosa), a compound synangium.     

SPORE MORPHOLOGY IN THE CYATHEACEAE. I. THE PERINE AND SPORANGIAL CAPACITY: GENERAL CONSIDERATIONS

The literature on cyatheaceous spore morphology relative to the presence of a perine layer is reviewed, and evidence based on a sodium-hydroxide assay is presented indicating that the outer scultpine layer in certain cyatheaceous spores is perine. Perine so defined characterizes Metaxya, paleotropical and certain neotropical species of Sphaeropteris, nearly all species of Alsophila, all species of Nephelea, and certain species of Trichipteris and Cyathea. It is lacking in Lophosoria, many species of Trichipteris and Cyathea, and all species of Cnemidaria. Two major patterns of spore number per sporangium in the family are reported. Lophosoria, Sphaeropteris, Trichipteris, Cyathea, Cnemidaria, and probably Metaxya are characterized by 64-spored sporangia, whereas most species of Alsophila and all species of Nephelea are characterized by 16-spored sporangia. The congruence of this generic distribution of sporangial-capacity types with Tryon's phyletic arrangement of cyatheaceous genera supports the naturalness of his system. The intrasporangial germination of spores retained in dehisced and dispersed sporangia supports the suggestion that decreased spore number per sporangium in Alsophila and Nephelea may relate to the role of the sporangia as dispersal units. The decreased number of spores per sporangium is associated with a trend toward increase in the number of sporangia per sores, with the highest known count approaching 1000 sporangia per sorus. The Alsophila-Nephelea evolutionary line has probably not been ancestral in the phylogeny of the more advanced groups of ferns.     

THE VASCULAR SYSTEM OF DOLEROTHECA AND ITS PHYLOGENETIC SIGNIFICANCE

The course of the vascular system in the proximal end of Dolerotheca formosa is described. Vascular bundles flare outward and downward immediately upon entering from the peduncle. These bundles are located in radiating septa just beneath the cover and give off small lateral bundles pinnately. Laterals from adjacent septal bundles meet, fuse, and extend downward in the parenchyma plate separating paired sporangia. The septal bundles, therefore, alternate in position with the parenchyma plate bundles and are interpreted as remnants of an ancestral bifurcating pinna system, which bore pendent sporangia along each side of supporting rachises. This interpretation differs from both the Codonotheca aggregation and plicated Whittleseya hypotheses recently advanced to explain the evolutionary pathway by which this complex pteridosperm pollen organ evolved.     

ON PHENOTYPIC EXPRESSION,MORPHOGENETIC PATTERN AND SYNANGIUM EVOLUTION IN PSILOTUM

Both the typical form and the appendageless variant of Psilotum nudum produce terminal synangia at the ultimate tips of the aerial axes. One clone in particular of the typical appendaged form produced synangia entirely at the tips of the aerial branches, as in the appendageless variant, and also developed occasional lateral transitional entities on the upper aerial axis displaying appendagelike and axislike morphological qualities. A developmental comparison of synangium development at the ultimate tips of aerial branches and of unusually elongating and normal sized fertile-appendages showed that the morphogenetic pattern of synangium development was similar. Anatomical and morphological evidence showed the synangium to be derived from terminal subdivisions or bifurcations of the apical meristem of each structure studied. This supports the phyletic concept that the synangium of the Psilotaceae is basically terminal to an axis or an axis homologue, and that it probably evolved from terminal bifurcative branching. Occasional multiple sporangium lobes may be formed on a P. nudum synangium which may not be represented by vascular bundles. Two hypothetical phyletic models of synangium evolution are proposed that could be used to explain this phenomenon and which should be tested by further evidence. Typical and appendageless P. nudum were compared in their morphogenetic pattern developed at the upper axis vegetative apical meristem, and a reconciliation was made between the structuring of the apparently disparate forms, which involved the presence or absence of serial ordering in apical derivatives. It is suggested that this could serve as a model for appendage evolution in the family Psilotaceae.     

A NEW LYGINOPTERID POLLEN ORGAN WITH ALVEOLATE POLLEN EXINES

A new lyginopterid pollen organ is described based upon specimens occurring in a single coal ball from the Providence, Kentucky locality. Seven to nine beaked sporangia are fused together at their proximal ends forming a common synangial chamber; synangia are joined together in clusters of two or three. In situ prepollen is similar to Cyclogranisporites and Verrucosisporites sporae dispersae. The thick exine has a lamellate nexine and a prominent alveolate sexine.     

THE SYNANGIATE NATURE OF DOLEROTHECA

Transverse sections through the distal region of a Dolerotheca specimen in a coal ball from the Calhoun coal of Illinois show the double rows of sporangia where they are free from one another. This leads to the conclusion that the sporangial rows should be regarded as extending inward from the campanulum cover rather than radiating outward from the center. This observation suggests that Dolerotheca is a synangium composed of many sporangial units of different lengths.     

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.     

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.     

AN ANATOMICALLY PRESERVED POTONIEA WITH IN SITU SPORES FROM THE PENNSYLVANIAN OF ILLINOIS

Specimens preserved in coal balls collected from the Herrin (No. 6) Coal in Illinois reveal the internal structure of the pteridosperm pollen organ long known as the compression fossil, Potoniea. Cylindrical sporangia are concentrically arranged in a campanulate body and the end of each sporangium projects freely from the distal face of the organ. Pollen is trilete and without a distal germinal aperture. The exine is structureless and has a separable, perine-like layer adorned with orbicules.     

A marattialean fern,Scolecopteris libera n. sp., from the Asselian (Permian) of Inner Mongolia,China

Scolecopteris libera n. sp. is established on partial three-dimensionally preserved materials containing both sterile and fertile fronds collected from the volcanic tuff at the top of the Taiyuan Formation at the Wuda Coalfield, Inner Mongolia, China. It is characterized by tripinnate fronds with rachises over 200 mm in diameter; and long lanceolate or falciform pinnules with thick veins. Pinnules are of the pecopterid-type, with those at the base of the ultimate pinnae usually divided into small lobes. Eight to twelve circular synangia are arranged in two rows along the midvein. Synangia are borne on a short pedicel attached to the middle of lateral veins. A synangium is composed of 7–10 fusiform exannulate sporangia with pointed apices. The sporangia are free among each other beyond the base of synangia. In situ microspores of the Cyclogranisporites leopoldii type differ from all other in situ Paleozoic marattialean spores.     

VALLITHECA VALENTIA GEN. ET SP. NOV., PERMINERALIZED SYNANGIA FROM THE MIDDLE PENNSYLVANIAN OF OKLAHOMA

Vallitheca valentia gen. et sp. nov. is a permineralized, synangiate fructification of probable seed fern affinities from the Wewoka Formation (Desmoinesian) near Ada, Oklahoma. Synangia are oblong to pyriform, average 1 cm long, and contain 32 to 48 tubular sporangia embedded proximally in ground tissue. The sporangia are arranged within the synangia in a unique pattern: a ring of peripheral sporangia dehisces toward the center of the synangium and a central group of sporangia dehisces outward. This unique arrangement is unlike that of any currently recognized group.