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.     

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.     

MILLAYA GEN. N., AN UPPER PALEOZOIC GENUS OF MARATTIALEAN SYNANGIA

Anatomically preserved fertile marattialean pinnules from the Lower Permian (Wolfcampian) Bursum Formation in Tularosa, New Mexico, and isolated synangia from the Upper Pennsylvanian (Missourian) Stanton Formation near Tyro, Kansas, are described. The sessile synangia are partially sunken into the pinnule lamina and arranged abaxially in a single row on each side of the midrib. Comparisons are made with other fossil and modern marattialeans and the evolutionary significance considered.     

Zhutheca Liu, Li et Hilton gen. nov., the fertile pinnules of Fascipteris densata Gu et Zhi and their significance in marattialean evolution

The morphology of the fertile pinnules of Fascipteris densata Gu et Zhi have been subjected to a detailed morphological investigation based on recently collected specimens from the Upper Permian of Yunnan Province, south China. These investigations have revealed that this species possesses synangia of the Asterotheca-type, situated in two or three rows either side of the midrib on a Fascipteris-type pinnule. This arrangement of Asterotheca-type synangia is extremely unusual considering that all other reports of this genus are with a single row of synangia located on each side of the midrib of a pecopteroid-type pinnule. As a consequence of this unique morphological arrangement, a new genus, Zhutheca densata Liu, Li et Hilton gen. et comb. nov. has been created to distinguish this material from other specimens of the Asterotheca and Fascipteris types. The structure and arrangement of the fertile pinnule of Zhutheca are compared with other Palaeozoic and Mesozoic marattialean taxa with which it shares certain features of its morphology. In addition, the stratigraphic, evolutionary and phylogenetic significance of Zhutheca are considered in detail.     

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.     

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.     

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.     

Reconstructing the Pennsylvanian-Age Filicalean Fern Botryopteris tridentata (Felix) Scott

Numerous anatomically preserved fragments of the Middle Pennsylvanian age filicalean fern, Botryopteris tridentata, occur in coal balls collected at the Pittsburgh and Midway Coal Company mine near Baxter Springs, Kansas. Included are the first fertile specimens of the species, evidence of complete vegetative frond architecture, and fronds that are specialized for vegetative propagation. Rhizomes are erect and unbranched, have helical phyllotaxis and short internodes, and typically display an ectophloic solenostele. Fronds are tripinnately compound with lobed pinnules that have open, dichotomous venation. Fertile pinnae or individual pinnules are interspersed among vegetative frond segments and produce sori of annulate sporangia beneath veins on the abaxial pinnule surface. Fertile pinnule lobes are rolled toward the abaxial surface to enclose the sori. Sporangia have a horizontally elongated biseriate annulus located near the short broad stalk and produce tetrahedral-shaped trilete spores with coarse spines. Epiphyllous branches diverge from the stipe or rachis, and some fronds produce only branches. This fern is reconstructed as having short stems. Helically arranged fronds are either pinnately dissected with lobed vegetative pinnules and abaxially rolled fertile pinnules or are specialized for vegetative propagation. The latter functioned as the foliar equivalent of stolons. While some characters of the B. tridentata plant are similar to those of Botryopteris forensis, generitype of the Botryopteridaceae, others are more comparable to those of Psalixochlaena cylindrica, generitype of Psalixochlaenaceae, suggesting the need for reevaluation of systematic relationships among species of the Botryopteridaceae and Psalixochlaenaceae.     

Cyathea cranhamii sp. nov. (Cyatheaceae), anatomically preserved tree fern sori from the Lower Cretaceous of Vancouver Island, British Columbia

Permineralized cyatheaceous sori occur among remains of conifers, fungi, and other plants in newly discovered calcareous concretions from Early Cretaceous (Barremian) marine sediments of Vancouver Island, British Columbia, Canada. Sori are superficially attached in two rows to narrow pinnules and display a globose sphaeropteroid indusium. Annulate sporangia with multicellular stalks diverge from a basal, vascularized receptacle. The nearly vertical uniseriate annulus is not interrupted by the stalk. The sporangia bear 64 trilete spores with perispore sculpturing that ranges from irregular granulate/echinate to prominent rodlets. These specimens, described as Cyathea cranhamii sp. nov., are the first anatomically preserved tree fern sori from the fossil record. They represent the most ancient evidence for fertile structures of the Cyatheaceae and demonstrate that essentially modern species of cyatheaceous tree ferns had evolved by the Early Cretaceous.     

Paralygodium meckertii sp. nov. (Schizaeaceae) from the Upper Cretaceous (Coniacian) of Vancouver Island, British Columbia, Canada

More than 50 specimens of permineralized fertile pinnules with abaxially borne sporangia have been discovered in calcareous marine nodules from the Upper Cretaceous (Coniacian) Comox Formation from the Eden Main localities on Vancouver Island, British Columbia, Canada. Isolated pinnules 1.6–3.0 mm wide × 1.6–2.8 mm long are lobed and abaxially enrolled to form irregular globose structures. Pyriform sporangia 216–300 μm wide × 360–468 μm long occur in two rows on the abaxial surface of pinnule lobes. Sporangia have an apical annulus of 15–18 cells. Spores are tetrahedral and trilete, 33–42 μm in diameter, with straight to concave interradial sides, laesurae extending nearly to the equator, and a psilate exine. Spores are assignable to the sporae dispersae genus Deltoidospora. Fertile pinnules are compared to fossils of Anemia poolensis and two previously described species of Paralygodium, and show closest similarities to P. vancouverensis from the Eocene of British Columbia. The Cretaceous Eden Main specimens differ in number of pinnule lobes and their morphology and are described as a new taxon: P. meckertii sp. nov. This discovery extends the Cretaceous geographic range of Paralygodium from Japan to North America and adds to our knowledge of the diversity of extinct schizaeaceous ferns.     

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.     

A new microsporangiate organ from the Lower Carboniferous of the Novgorod Region,Russia

A new species of the genus Telangiopsis, T. nonnae O. Orlova et Zavialova, was described on the basis of a microsporangiate organ from the Lower Carboniferous deposits of the Novgorod Region. The morphology of branching fertile axes, synangia, and sporangia was thoroughly studied. The three-dimensional system of fertile axes branches monopodially; ultimate axes bear numerous connivent bunches of synangia, which consist of three to six basally fused elongated ovate sporangia. The morphology and ultrastructure of prepollen grains were studied, which were extracted from the rock matrix surrounding the sporangia. The two-layered exine includes a well-developed endexine and an alveolate ectexine, with one-three rows of large thin-walled alveolae. The new species was compared with other Early Carboniferous microsporangiate organs.     

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.     

Anatomically preserved Woodwardia virginica (Blechnaceae) and a new filicalean fern from the middle Miocene Yakima Canyon flora of central Washington, USA

Anatomically preserved Woodwardia virginica (Blechnaceae) and a newly recognized onocleoid fern are described from the middle Miocene Yakima Canyon flora of central Washington State, USA. Identification of the W. virginica fossils is based on a combination of vegetative pinnules, rhizome and stipe anatomy, and fertile pinnules with indusiate sori and sporangia like those of extant W. virginica. Fronds are isomorphic. Vegetative pinnae are elongated and pinnatifid, with a secondary vein paralleling the midvein. Secondary veins of the pinnule lobe anastomose to form primary areoles and are either simple or dichotomize toward the margin. Rhizomes have a simple dictyostele with 3-5 cauline vascular bundles and often a sclerotic hypodermis. Leaf traces contain two large adaxial vascular bundles that occur laterally and adaxially, flanking an arc of 4-6 smaller bundles. Fertile pinnules have linear sori that are somewhat embedded in the laminae and are enclosed by a thin indusium. Leptosporangia display a vertical annulus and an elongated stalk. A second fern, Wessiea yakimaensis gen. et sp. nov., is represented by anatomically preserved branching rhizomes and attached frond bases that conform to the Onoclea-type pattern of rhizome and frond-base vasculature. Rhizomes have a simple dictyostele of 4-5 cauline meristeles. Leaf divergence is helical, with paired hippocampiform rachial traces. These two ferns occur in the same matrix with specimens of Osmunda wehrii. They demonstrate that filicalean fern assemblages similar to those of extant temperate floras were well established in western North America by the middle Miocene and further emphasize the exceptional species longevity of some homosporous pteridophytes.     

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.     

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.     

A new species of the marattialean fern Scolecopteris (Zenker) Millay from the uppermost Permian of Guizhou Province, south-western China

Several isolated marattialean synangia and sporangia are reported from coal balls collected from Coal Seam No.1 (C605) in the uppermost Permian Wangjiazhai Formation in Guizhou Province, south-western China. The synangia are radially symmetrical with diameters between 0.8 and 1.2 mm and are 1.7 mm long, consisting of 3–4 elongate sporangia that are fused basally, free distally and possess a pointed apex. The outer-facing sporangial wall is 4–5 cells thick and conspicuously differentiated. Spores are trilete, have a granular ornamentation and are nearly round equatorially with a diameter of 55–60 µm. Comparisons with other anatomically preserved Palaeozoic marattialean synangia from the Euramerican and Cathaysian floras permit their assignment to the genus of Scolecopteris (Zenker) Millay. In this species the thick, outer-facing sporangial walls and large trilete spores are features consistent with those of the Oliveri Group within Scolecopteris , a group that has previously been considered primitive within this genus. Distinctions from all other previously recognized species within the Oliveri Group lead to the creation of a new species, S. guizhouensis sp. nov. This species is the youngest of the reported species of Scolecopteris recognized from the Euramerican and Cathaysian floras, and provides important evidence on the organization of marattialean ferns from the Upper Permian strata of south China.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 279–288.     

A new late Westphalian fossil marattialean fern from Nova Scotia

Sydneia manleyi gen. et sp. nov. is based on part of a fertile frond from the upper Westphalian D of the Sydney Coalfield, Nova Scotia, Canada. It has small synangia composed of laterally fused sporangia that are elongate and with a circular cross-section. The sporangia yielded variably sized monolete and trilete spores with laevigate and microspinate ornamentation; intermediate forms were also observed. The spores can be correlated with the sporae dispersae species Latosporites minutus , Punctatosporites oculus and Laevigatosporites minimus . Size distribution of the spores is variable and highly skewed, suggesting heterogeneity of the spores within the sporangium. Spore ultrastructure indicates that the fossil is part of a fern, and the morphology of the spores and synangia indicate marattialean affinities.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 199–212.     

Ashicaulis woolfei n. sp.: additional evidence for the antiquity of osmundaceous ferns from the Triassic of Antarctica

Numerous small fern trunks and dispersed osmundaceous frond fragments occur within a Middle Triassic silicified peat near Fremouw Peak in the Transantarctic Mountains of Antarctica. These specimens form the basis of a new species of osmundaceous ferns that further helps to characterize the early Mesozoic vegetation of high latitude Gondwana. Ashicaulis woolfei n. sp. consists of small, upright trunks with a persistent armor of frond bases, adventitious roots, and vegetative frond parts. In cross section the trunks are ～2.5 cm in diameter and include up to 45 frond bases. Stems range from 5 to 8 mm in diameter with a xylem cylinder of 8-9 xylem segments separated by leaf gaps. Phyllotaxy is variable, approaching 2/5 or 3/8, with 10-12 frond traces in the cortex. Stipes have parenchymatous, stipular wings that are usually devoid of sclerenchyma; fronds are pinnate with alternate-subopposite pinnatifid pinnules. Although the absence of fertile pinnules and sporangia precludes assigning the fossils to a living genus, this species demonstrates that ferns with stelar architecture and histology similar to Osmunda subgenus Osmundastrum (Osmundaceae) were present in the Southern Hemisphere by the mid-Triassic.