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.     

A WOODY LYCOPSID STEM FROM THE NEW ALBANY SHALE (LOWER MISSISSIPPIAN) OF KENTUCKY

A lycopsid axis from the New Albany Shale (Sanderson Formation) of Kentucky is described. The stem, which branches dichotomously, is 45 mm in diameter and is characterized by a relatively narrow parenchymatized protostele, a 3.0 mm-thick cylinder of secondary xylem, a tripartite cortex, and a periderm that is more than 5.0 mm thick. The secondary xylem is composed of uniseriate and biseriate vascular rays and narrow tracheids with scalariform wall thickenings on both radial and tangential walls. The periderm is characterized by elongate, thick-walled cells, some of which broaden tangentially in the outer part of the tissue forming zones that appear wedge-shaped in cross section. Surface features of the axis, including leaf bases, are not preserved. The stem is tentatively regarded as a member of the Lepidodendrales in accordance with the numerous anatomical characters that it shares with more recent representatives of the order. Because the external morphology is not known, however, the possibility exists that the axis corresponds to a protolepidodendralean taxon currently known only from compression and/or impression remains or some other nonlepidodendralean plant that produced secondary xylem. The extremely narrow profile of the secondary xylem tracheids (relative to other arborescent lycopsids) is interpreted as evidence that the plants grew in a habitat that was substantially drier than the Upper Carboniferous coal swamps.     

On Paleozoic plants from marine strata: Trivena arkansana (Lyginopteridaceae) gen. et sp. nov., a lyginopterid from the Fayetteville Formation (middle Chesterian/Upper Mississippian) of Arkansas, USA

Five permineralized seed fern stems from the Fayetteville Formation (middle Chesterian/Upper Mississippian) of Arkansas conform to the concept of lyginopterid seed ferns. However, these specimens are unlike all previously reported lyginopterids, and the name Trivena arkansana (Lyginopteridaceae) gen. et sp. nov. is proposed. The stems are up to 30 by 19 mm in diameter and have pentagonal pith and eustele of five cryptic sympodia. Secondary tissues include abundant xylem with numerous wide rays and phloem surrounded by a periderm. The cortex is parenchymatous with abundant sclerotic clusters: some clusters are randomly dispersed and some are in discontinuous rows. Sclerenchyma bands form the "Dictyoxylon"-type outer cortex. Leaf traces diverge in a 2/5 phyllotaxy. Traces, accompanied by concentric secondary xylem, increase in size as they extend through the secondary xylem of the stem. The trace assumes a squat C shape at the outer margin of the secondary xylem and in the cortex divides into three discrete bundles, each surrounded by secondary xylem. Galleries within the phloem contain arthropod coprolites and exhibit wound response, suggesting plant-arthropod coevolution. The discovery of this new lyginopterid stem adds to the growing list of unique taxa described from the Fayetteville Formation and further solidifies its reputation as one of the most important Upper Mississippian plant fossil sites in North America.     

QUAESTORA AMPLECTA GEN. ET SP. N., A STRUCTURALLY SIMPLE MEDULLOSAN STEM FROM THE UPPER MISSISSIPPIAN OF ARKANSAS

A well preserved, permineralized seed fern stem is described from the Upper Mississippian Fayetteville Formation of north central Arkansas. Quaestora amplecta gen. et sp. n. is 41.6 cm long and exhibits six pairs of decussate, highly decurrent petiole bases. The stem has a cruciform, exarch protostele with prominent secondary xylem, vascular cambium and secondary phloem. Leaf traces are terete and occur as an outer ring with a small number of internal strands. The cauline vasculature, leaf-trace production, petiolar anatomy and several other features indicate that this specimen represents the most structurally simple and geologically ancient medullosan stem presently recognized.     

A new genus of filicalean fern from the Lower Permian of China

Recent investigation on the Cathaysian flora of the Lower Permian Taiyuan Formation in Hebei Province, North China has led to the discovery of a new type of fern, Rastropteris pingquanensis gen. et sp. nov. Preserved as a permineralization, the stem, with a mantle of petiole bases and roots, shows a unique combination of anatomical characters. The vascular strand consists of a solid protostele with uniformly elongated tracheids and mesarch maturation of the xylem. In transverse section, the configuration of leaf trace xylem changes from reniform endarch to a tangentially elongated strand adaxially recurved at each end with several adaxial ridges. Stem and petiole cortex contains abundant sclerotic tissue with an interstitial tissue developed within the mantle of petiole bases and roots. A reinvestigation of Grammatopteris rigollotii , from the Permian of France, the type species of the genus allows interpretation of the latter as closer to Rastropteris than any other fern. Both taxa are provisionally reported to an incertae sedis group of filicalean ferns possibly involved in the origin of Osmundaceae.     

CAULINE VASCULATURE AND LEAF TRACE PRODUCTION IN MEDULLOSAN PTERIDOSPERMS

Medullosa and Sutcliffia specimens from the Paleozoic of North America and Europe are examined to determine the architecture of the cauline vasculature and mode of leaf trace production. Emphasis is placed on the identification and characterization of protoxylem strands and their relationship to leaf trace production. Organization of the primary xylem varies from a single protostele to a dissected stele composed of two to many more or less independent bundles. In Medullosa the bundles of primary xylem are each surrounded by secondary xylem, forming separate segments of vascular tissue (‘steles’ of previous workers). These vascular segments may divide and fuse at different levels in the stem. A definite number of protoxylem strands occur near the periphery of the primary xylem. The protoxylem strands divide at intervals producing protoxylem to the departing leaf traces. Leaf traces thus formed arise from all the vascular segments in a coordinated and predictable way and pass outward through emission areas in the secondary xylem. This type of cauline vascular architecture is compared to that of other seed plants. The vascular system of Medullosa stems is interpreted as a dissected monostele. Sympodial vascular architecture has apparently evolved from a protostele separately within the medullosan pteridosperms.     

PROTEOKALON A NEW GENUS OF PROGYMNOSPERMS FROM THE DEVONIAN OF NEW YORK STATE AND ITS BEARING ON PHYLOGENETIC TRENDS IN THE GROUP

Proteokalon gen. nov. is described from the Upper Devonian Catskill deposits of New York. Two orders of branching and ultimate appendages are preserved' by petrifaction and by compression. The first order bears branches decussately and has a skewed four-armed protostele that occasionally dichotomizes. Second-order branches dichotomize rarely and most have T-shaped or three-armed protosteles. They bear ultimate appendages alternately, either in lateral pairs, or singly from the abaxial side. These appendages divide several times in one plane. Their vascular strand is terete. Maturation of the primary xylem is mesarch, and it consists of tracheids and parenchyma. Secondary xylem and phloem and a periderm are present. The outer cortex has a system of hypodermal fibers. Proteokalon is most similar to Tetraxylopteris and Triloboxylon of the Aneurophytales. A comparison of the stratigraphic occurrence of Protopteridium, Aneurophyton, Tetraxylopterism, Sphenoxylon, Triloboxylon, and Proteokalon suggests some evolutionary trends among the Aneurophytales.     

STENOKOLEOS BIFIDUS SP. N. IN THE UPPER DEVONIAN OF NEW YORK STATE

Stenokoleos is a genus for petrified axes from the Mississippian New Albany Shale to which an Upper Devonian occurrence in New York is added. Two orders of branching were known and the plant was thought to be related to coenopterid ferns. The new petrified axes from New York reveal three orders of branching. A pair of rachides emerges from one side of the stem at each node. Their position alternates at successive nodes (distichous). Each rachis bears alternately arranged pinnae. The shape of the xylem strand and the number of protoxylem areas are variable. Traces to the pairs of rachides arise either as two separate strands or as a single strand that is presumed to divide while still within the cortex of the stem. Traces to pinnae are ellipsoid or clepsydroid. Tracheids are scalariform and uni- or biseriate, circular-bordered pitted. Peripheral loops are present in all orders of branches. Protoxylem strands are numerous and maturation is mesarch. Cortex is parenchymatous where it is preserved but outer cortex is missing. Stenokoleos and Reimanniopsis are placed in a new family, Stenokoleaceae. This is classified as Incertae Sedis among Pterophytina in Tracheophyta. It is suggested that the plant is related more closely to the Mississippian pteridosperms Tristichia and Tetrastichia than to the coenopterid ferns.     

扁圆封印木（相似种）茎干的解剖特征

贵州省水城矿区晚二叠世煤核中扁圆封印木（相似种Ｓｉｇｉｌｌａｒｉａ ｃｆ．ｂｒａｒｄｉｉＢｒｏｎｇｎ．）茎干的主要解剖特征如下：管状中柱,具多边形薄壁细胞组成的髓。初生木质部成环带状,外缘呈规则的齿槽状,向心式发育。次生木质部显束状特征,横切面管胞为方圆至长方形,纵切面为梯状壁增厚,并具流苏纹。射线１—２列细胞宽,数个至十余个细胞高。叶迹起源于初生木质部外缘的槽中,中始式,但以向心发育为主。     

STRUCTURALLY PRESERVED FOSSIL PLANTS FROM ANTARCTICA. I. ANTARCTICYCAS,GEN. NOV., A TRIASSIC CYCAD STEM FROM THE BEARDMORE GLACIER AREA

Silicified stems with typical cycadalean anatomy are described from specimens collected from the Fremouw Formation (Triassic) in the Transantarctic Mountains of Antarctica. Axes are slender with a large parenchymatous pith and cortex separated by a narrow ring of vascular tissue. Mucilage canals are present in both pith and cortex. Vascular tissue consists of endarch primary xylem, a narrow band of secondary xylem tracheids, cambial zone, and region of secondary phloem. Vascular bundles contain uni- to triseriate rays with larger rays up to 2 mm wide separating the individual bundles. Pitting on primary xylem elements ranges from helical to scalariform; secondary xylem tracheids exhibit alternate circular bordered pits. Traces, often accompanied by a mucilage canal, extend out through the large rays into the cortex where some assume a girdling configuration. A zone of periderm is present at the periphery of the stem. Large and small roots are attached to the stem and are conspicuous in the surrounding matrix. The anatomy of the Antarctic cycad is compared with that of other fossil and extant cycadalean stems.     

Anatomical Characteristics of the Stem of Sigillaria cf. brardii from Coal Balls in Guizhou Province,China

The stem specimens of Sigillaria cf. brardii were collected from the coal balls of Upper Permian in Shuicheng Coal Mines in Guizhou Province. The main anatomical characteristics of Sigillaria cf. brardii are described as follows: The stem is siphonostelic, with pith composed entirely of polygonal parenchyma cells, there are secondary walls in some pith cell cavities these secondary walls show the characters of cell division. Surrounding the pith is the continuous cylindrical primary xylem which consists entirely of tracheids. The outermost, and part are the protoxylem elements show spiral secondary thickenings. In cross section, the outer edge of exarch primary xylem appears regularly sinuous, with trace of mesarch leaf originating from the furrows. The centripetal metaxylem is characterized by scalariform wall thickenings on the tracheids, and delicated strands of secondary wall materials extending between abjacent bars, these structures are called fimbris, or williamson striations, and are characteristic in lepidodendrids. The secondary xylem consists of tracheids and vascular rays. The tracheids, too, have scalariform wall thickenings and fimbris. The rays are one-to twocell width and several to more than ten cells in height.     

Palaeosmunda Emended and Two New Species

The genus Palaeosmunda was established by R. E. Gould in 1970 based upon some Late Permian Osmundaceous trunks with well-developed leaf gaps and rhomboidal sclerotic ring within petiolar base seen in cross section. As he thinks that the latter character is more important than the former, this genus could not be assigned to any subfamily of Osmundaceae. However, the leaf gap is one of the most important characters in the structure of the fern stem, so the author suggests that this genus should be assigned to subfamily Osmundoideae and its diagnosis must be emended as follows: The genus Palaeosmunda is represented by some rhizomes (or trunks), roots and leaf bases of ferns which structurally are preserved, resembling Osmundacaulis but which can’t be assigned to any group of this genus. Stem containing an ectophloic dictyoxylic siphonostele; if tracheids present in the pith, they being multiseriate scalariform pitted; pith or cortex sometimes contain ing groups of secretory cells or sclerenchyma; number of leaf traces seen in a tran sverse section of cortex more than 30; leaf traces adaxially curvature, rarely oblong shaped; petiolar bases with or without stipular expansion, containing a C-shaped vascular strand; root diarch. Type species——Palaeosmunda williamsii. According to this diagnosis some primitive osmundaceous species with the leaf gaps, which have already found in Upper Permian and Lower Triassic, could be assigned to this genus. Two of them are P. williamsii Gould and P. playfordii Gould, and Osmundacaulis beardmorensis, which was from Lower Triassic of Antarctica in 1978, should be assigned to the genus Palaeosmunda. In this paper two osmundaceous new species: P. primitiva and P. plenasioides were found in the coal balls of Upper Permian age from Wangjiazhai of Shuicheng of Guizhou Province, China. P. primitiva is represented by two trunks; stem about 4 cm in diameter; stele actophloic dictyoxylic siphonostele; pith cavity about 3—4 mm in diameter, contianing parenchyma and tracheids; xylem cylinder thin, less than 10 tracheids in radial thickness, dissected by leaf gaps. Inner cortex about 1.5 cm thick, mainly parenchymatous, but sometimes containing a few sclerenchymatous; number of leaf traces seen in a transverse section about 50—60; leaf traces departing at 35—45º,open C-shaped at point of departure, gradually becoming shallow C-shaped or V-shaped in different parts; protoxylem in base of leaf traces single, endarch; when leaf traces pass through inner cortex, protoxylem biturcating. Petiole bases without stipular expansion, probablyloosely embracing the stem; xylem strand of potiole trace shallow C-shaped, surrounded by selerenchyma; sclerotic ring round, connected with single sclerenchyma mass in the concavity of the petiole trace. Root arising singly from leaf trace, diarch, with inner and outer cortex. P. plenasioides is represented by a rhizome; stem more than 4 cm in diameter; stele actophloic dictyoxylic siphonostele; xylem cylider with about 20 tracheids in radial thickness, dissected by leaf gaps; xylem bundle U-, O-, or crosier- (i.e. query-) shaped; pith and inner cortex parenchymatous, with many groups of secretory cells; leaf trace C-shaped, its base containing two endarch protoxylem groups; root diareh,with inner and outer cortex, arising singly from leaf trace or its base.     

The Givetian flora from Cairo, New York: Rhacophyton, Triloboxylon and Cladoxylon

Petrified Rhacophyton, Triloboxylon and Cladoxylon are described from the Givetian of eastern New York State. In cross section, specimens referable to Rhacophyton ceratangium have a mesarch primary xylem strand in the shape of a bar with swollen endS. Pycnoxylic secondary xylem surrounds the primary xylem. Vascular strands, interpreted as traces, are also present. Triloboxylon has a three-fluted primary xylem strand surrounded by secondary xylem. Several mesarch protoxylem areas are present in cross sectional view. The specimens of Cladoxylon , the first of this genus in the Middle Devonian of North America, show the typical polystelic pattern in cross section. Obvious secondary xylem and peripheral loops are absent. Lateral appendages were observed on two of the specimenS. A comparison of the Cairo and Gilboa floras indicates that they represent different ecological niches during the late Middle Devonian.     

OSMUNDA WEHRII,A NEW SPECIES BASED ON PETRIFIED RHIZOMES FROM THE MIOCENE OF WASHINGTON

Silicified rhizomes from Miocene strata near Yakima, Washington represent a new species of Osmunda. The stems are 8–13 mm in diameter and are surrounded by a thick sheath of adherent leaf bases, each of which shows stipular expansions typical of the Osmundaceae. The new species has an ectophloic siphonostele in which the xylem cylinder is dissected by leaf gaps with 12–14 strands being visible in a given stem cross section. Such sections also show 12–16 leaf traces in the cortex. The xylem of each leaf trace diverges from the xylem cylinder of the stem as an adaxially concave strand with its protoxylem organized into a single medial adaxial cluster. Initial bifurcation of the leaf-trace protexylem occurs as the leaf trace passes through the outer cortex of the stem. In the basal part of the stipular region of the petiole base, thick-walled fibers form an arch on the abaxial side of the sclerenchyma ring around the petiolar bundle. This arch persists throughout most of the length of the stipular region, with the thick-walled fibers becoming reorganized into two lateral masses in the distal part of the stipular region. Similar thick-walled fibers form an elongate strip of tissue in each wing of the stipule along with several small clusters scattered near the sclerenchyma ring. The new species belongs to the subgenus Osmunda and shows that during the Neogene, the latter existed as a group of closely related species much as it does today. Furthermore, Osmunda wehrii combines features of the modern O. regalis, O. japonica, and O. lancea with those of O. claytoniana and thus supports the inclusion of the latter species in the subgenus Osmunda.     

Anatomical Study of the Rachis of Compsopteris elliptica and the Systematic Position of Compsopteris

The anatomical characters of the rachis of Compsopteris elliptica ex Yang et Chen are described from the calc-petrified specimens of the Late Permian of Panxing, Guizhou, China. The main characters of its rachis are as follows (Plate I, 1–4; 6–9): The vascular bundles of 2– 2.5 cycles and ectophloic type. Phloem, consisting of 1–2 layers of cells. Protoxylem, composed of several small tracheids which are less than 20μm in diameter. Metaxylem, consisting of 2–3 layers of tracheids, about 30μm in diameter, scalariform thickenning. The sclerenchyma zone lies between the vascular bundles, its cell, small, 20–25 μm in diameter, some containing brown substances. The cortex could be divided into two zones: outer zone, consisting of thin-walled cells, and inner zone, thicken-walled cells, with secretory cavities. Epidermis, one layer, rectangular, some with contents. Compsopteris sp. (Plate I,5) 3–4 cycles vascular bundles, the same as C. elliptica in many respects; only larger than C. ellipptica in diameter. It may represent the base part of the rachis or larger rachis. Based on the shape of vascular bundles and the structure of xylem and phloem, Compsopteris is very similar to Angiopteris, Danaea etc., which all belong to the Marattiales. In addition, Huang et al. (1989) found that the sporangia of Compsopteris is similar to that of Danaeites, which belongs to the Maratriales. Therefore, Compsopteris doer not belong to the seed fern, bur the Marattiales.     

Sur les fougères Anachoropteris et Tubicaulis du Stéphanienfrançais. Description de Tubicaulis grandeuryi nov. sp.

New information is presented on the morphology of leaves and stems of the ferns Anachoropteris and Tubicaulis preserved in silicified material from the Stephanian of Grand'Croix (France). A rachis of Anachoropteris involuta bears laterally, in association with a pinna, an epiphyllous stem, with a vitalized protostele. This stem bears nine petioles and is almost identical in anatomy to the american species, Tubicaulis stewartii. A hand-ground section from the Grand'Eury collection, Paris, is described as Tubicaulis sp. and identified as a more distal region of a similar stem. Tubicaulis grandeuryi nov. sp. is another type, with haplostelic stem and petioles bearing pinna traces near their base. Axillary branching is described in this genus for the first time. The problem of redefining the systematic position of Tubicaulis and its relationship to Anachoropteris is discussed.     

PINUS WOLFEI,A NEW PETRIFIED CONE FROM THE EOCENE OF WASHINGTON

A silicified cone from the Late Eocene of Washington is described as a new fossil species of Pinus. The cone was probably 9–10 cm long and 3–5 cm at its widest diam in the living condition and is peculiar in having abundant resin canals in the secondary xylem of the axis arranged in three concentric rings near the cone base. The bract of the fossil is also unusual in having resin canals of distinctly unequal sizes and a vascular strand that is adaxially concave. In the absence of external features of the scale tips, these anatomical conditions along with the construction of the outer cortex of the axis of thick-walled cells suggest closest affinity of the new species with the subsections Contortae, Oocarpae, and Sylvestres of the section Pinus, subgenus Pinus.     

Todea from the Lower Cretaceous of western North America: implications for the phylogeny, systematics, and evolution of modern Osmundaceae

The first fossil evidence for the fern genus Todea has been recovered from the Lower Cretaceous of British Columbia, Canada, providing paleontological data to strengthen hypotheses regarding patterns of evolution and phylogeny within Osmundaceae. The fossil consists of a branching rhizome, adventitious roots, and leaf bases. The dictyoxylic stem has up to eight xylem bundles around a sclerenchymatous pith. Leaf traces diverge from cauline bundles in a typical osmundaceous pattern and leaf bases display a sheath of sclerenchyma around a C-shaped xylem trace with 2-8 protoxylem strands. Within the adaxial concavity of each leaf trace, a single sclerenchyma bundle becomes C-shaped as it enters the cortex. The sclerotic cortex is heterogeneous with an indistinct outer margin. The discovery of Todea tidwellii sp. nov. reveals that the genus Todea evolved by the Lower Cretaceous. A phylogenetic analysis combining morphological characters of living and extinct species with a previously published nucleotide sequence matrix confirms the taxonomic placement of T. tidwellii. Results also support the hypothesis that Osmunda s.l. represents a paraphyletic assemblage and that living species be segregated into two genera, Osmunda and Osmundastrum. Fossil evidence confirms that Osmundaceae originated in the Southern Hemisphere during the Permian, underwent rapid diversification, and species extended around the world during the Triassic. Crown group Osmundaceae originated by the Late Triassic, with living species appearing by the Late Cretaceous.     

CRENATICAULIS,A NEW GENUS OF DEVONIAN PLANTS ALLIED TO ZOSTEROPHYLLUM,AND ITS BEARING ON THE CLASSIFICATION OF EARLY LAND PLANTS

The name Crenaticaulis verruculosus is proposed for slender, pseudomonopodially and dichoto-mously branching plants that bore opposite to subopposite sporangia along the stem and two rows of prominent, multicellular teeth on opposite sides of the terete axis. Epidermal cells were either narrow and elongate parallel to the stem or short with a papillate outer tangential wall. Axillary tubercles were present on one side of the stem near lateral branches. Some tubercles bore remnants of branches. Sporangia dehisced along their distal margins into two unequal halves. No spores were found. Occasional short lengths of stem were petrified by iron pyrite. Sections revealed a cortex consisting of four to six rows of thick-walled cells and a xylem strand. The strand was elliptical in transverse section and maturation was exarch. Tracheids were chiefly scalariform. It is suggested that the so-called axillary tubercles, known in several plants of Devonian age, were scars of rhizophores like those in the modern genus Selaginella. The plant is referred to the subdivision Zosterophyllophytina.