IBYKA AMPHIKOMA,GEN. ET SP. N., A NEW PROTOARTICULATE PRECURSOR FROM THE LATE MIDDLE DEVONIAN OF NEW YORK STATE

Ibyka gen. n. is described from late Middle Devonian compressions and petrifactions collected in eastern New York State. It is a robust plant of which three orders of branching and ultimate appendages (leaves) are known. The latter dichotomize up to five times, are arranged spirally on all orders of branching, are three-dimensional, and all orders are terete in cross section. Fertile appendages, homologous with the sterile, are terminated by sporangia. The protostele has five or six arms, the maturation is mesarch, and the protoxylem disintegrates leaving lacunae at the tips of the arms. Traces to appendages are terete and arise spirally from the tips of the arms. The primary xylem consists of tracheids only, the phloem of thin-walled cells and probable tanniniferous cells. The cortex consists of parenchyma and groups of sclereids. Secondary xylem is lacking. Ibyka is placed in a new order, Ibykales, close to the Hyeniales (protoarticulates) and to the Pseudosporochnales all three of which probably evolved from Trimerophytina.     

Rotafolia songziensis gen. et comb. nov., a sphenopsid from the Late Devonian of Hubei, China

A sphenopsid from the Upper Devonian (Famennian) Xiejingsi Formation, south-western Hubei Province, China, previously named as various species in Sphenophyllum , Hamatophyton , Bowmanites and Sphenophyllostachys , is now reinvestigated and assigned to a new taxon, Rotafolia songziensis gen. et comb. nov. Its ribbed axes are anisotomous and possess slightly expanded nodes. Lateral axes are inserted at nodes on main axes. Whorls of much divided vegetative leaves are attached at nearly right angles to nodes of basal axes, and at acute angles to nodes of terminal axes. There are six leaves per whorl. The terminal strobilus includes a central axis and verticils of fertile units. Each fertile unit consists of a bract and numerous sporangia. The margin of the elongate-cuneate bract bears a distal and many lateral elongate segments. Clusters of elongate sporangia are abaxially attached to the base of the bract at the same level. The axis has an actinostele, composed of a three-ribbed, exarch primary xylem and radial secondary xylem. Although Rotafolia songziensis closely resembles Hamatophyton verticillatum in axis character, leaf morphology and primary xylem type, they are quite different in strobilar structure. Taxonomically, Rotafolia is placed in the order Sphenophyllales by three well-defined characters: 1) whorled appendages; 2) ribbed protosteles; 3) exarch primary xylem maturation.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 21–37.     

A FERTILE AXIS OF TRILOBOXYLON ASHLANDICUM,A PROGYMNOSPERM FROM THE UPPER DEVONIAN OF NEW YORK

A partially petrified impression of Triloboxylon ashlandicum (Aneurophytales) is the first recognized fertile axis of the genus. Identification of the fertile axis rests on the similarity of its anatomy with that of previously described vegetative specimens. Fertile organs replace some vegetative branches along part of the main axis. Fertile organs are twice dichotomized in one plane and bear elongate sporangia arranged pinnately. Vegetative branches differ in that they bear the ultimate appendages of the plant helically. The latter organs dichotomize many times in one plane. Although similar in size and morphology to the ultimate appendages, the fertile organs are homologous by position and vascular supply to the vegetative branches which they replace. Sporangia of T. ashlandicum dehisce longitudinally and terminate in an apiculate tip. Spores are unknown. Fertile organs of T. ashlandicum resemble those of other Aneurophytales and support the earlier placement of Triloboxylon in the order on anatomical grounds. T. ashlandicum differs from other Aneurophytales, however, by bearing vegetative organs at the distal end of its fertile axis.     

Clevelandodendron ohioensis,gen. et sp. nov., a slender upright lycopsid from the late Devonian Cleveland Shale of Ohio

Clevelandodendron ohioensis Chitaley & Pigg gen. et sp. nov. is an almost entire lycopsid plant known from a single compressed specimen from the Cleveland Shale member of the Upper Devonian Ohio Shale. This unique specimen is 125 cm long, consisting of an unbranched, slender, monopodial axis with a partially preserved plant base bearing thick appendages at one end, and a compact, terminal ovoid bisporangiate strobilus at the other. The stem is 2 cm wide for most of its length. Visible on the decorticated stem surface are helically arranged, elongate leaf traces and laterally compressed, slender leaves along the stem margin. The plant base bears 4-6 thick appendages. The terminal strobilus is compact, ovoid, 9 cm long and up to 6 cm wide, morphologically similar to those of some Lepidodendrales, and bears helically arranged sporophyll/sporangium complexes with narrow bases and distal laminae up to 18 mm long, turned upward. Megaspores are 320-360 μm, trilete and laevigate, lacking a gula; microspores are 30-42 μm, trilete, indistinctly punctate and possibly assignable to Calamospora or Punctatisporites. Clevelandodendron demonstrates that slender unbranched lycopsids with an isoetalean plant habit similar to the Carboniferous genera Chaloneria and Sporangiostrobus and Triassic Pleuromeia-like forms were present as early as the Late Devonian. The early occurrence of this unique habit suggests that diversification within the isoetalean clade sensu Rothwell and Erwin (including both Isoetales and Lepidodendrales) was well established prior to the Carboniferous.     

THE PRIMARY VASCULATURE OF CORDAIANTHUS CONCINNUS

The pollen strobilus Cordaianthus concinnus is examined as a possible indicator of the basic pattern of vascular architecture in stems of the Cordaitales. Bract traces arise from two points in the stele of the bilateral primary axis and diverge to the regularly arranged, four-ranked bracts. Tracheids to the axillary secondary shoots arise as two traces that flank the position of bract trace emission. Distally, the secondary shoot traces unite to form a stele that becomes increasingly dissected at successively higher levels. Although radially aligned, these tracheids show thickening patterns on all walls and are not separated by vascular rays; they are therefore interpreted as primary xylem. The traces form sympodia that are similar to those of typical eustelic gymnosperms. Scale traces from the secondary shoots arise by the tangential division of an individual axial bundle and occur in arrangements that range from a ½ to a % spiral. The vascular architecture of these secondary axes is interpreted as the equivalent of that in the stems of extant conifers with spiral phyllotaxis.     

WOOD ANATOMY OF BUBBIA (WINTERACEAE), WITH COMMENTS ON ORIGIN OF VESSELS IN DICOTYLEDONS

Quantitative and qualitative features of wood anatomy are reported for ten collections of seven species of Bubbia. Variations on the basic plan for Winteraceae can be interpreted in terms of taxonomic and ecological distinctions. Tracheid length is correlated with plant size and habit: tracheids are shortest in shrubs. Tracheid wall thickness and ray cell wall thickness distinguish species. Ray cell procumbency and multiseriate ray width increase with age. Growth rings occur only in a species from stream margins. SEM studies reveal absence of a warty layer within tracheids. Helical thickenings are absent. Presence of these two features in Pseudowintera may be correlated with the cool temperate habitats of that genus. Overlap areas of tracheids in Bubbia show various degrees of scalariform pitting, ranging from none (B. semecarpoides) to abundant presence (B. balansae). Perforation-like pits in tracheids of the latter prove, with SEM studies, to have pit membranes containing porosities less than 1 μm in diameter. Scalariform pitting on overlap areas is absent in earlier secondary xylem and increases during later secondary xylem. Scalariform lateral wall pitting can occur in abnormally wide tracheids formed after pauses in cambial activity. These facts show that primitive dicotyledon woods like those of Bubbia can activate genetic information for scalariform end wall patterns and lateral wall pitting such as primitive vessels show without the intervention of paedomorphosis. Paedomorphosis in dicotyledon woods is held still to apply only to special herbaceous and herblike growth forms, not to primarily woody plants. Progenesis (in xylem, loss of secondary xylem) is not held to be necessary to account for the scalariform patterns seen in tracheary elements of primitive dicotyledons. Reasons are given for rejection of the hypothesis that Winteraceae and other woody dicotyledons (Amborella, Sarcandra, Tetracentron, Trochodendron) are secondarily vesselless.     

Changes in the arrangement of cellulose microfibrils associated with the cessation of cell expansion in tracheids

 The relationship between the cessation of cell expansion and formation of the secondary wall was investigated in the early-wood tracheids of Abies sachalinensis Masters by image analysis and field emission scanning electron microscopy. The area of the lumen and the length of the perimeter of the lumen of differentiating tracheids increased from the cambium towards the xylem. These increases had just ceased in the case of tracheids closest to the cambium in which birefringence was first detected by observations with a polarizing light microscope. Cellulose microfibrils (MFs) deposited on the innermost surfaces of radial walls were not well ordered during the expansion of cells, but well ordered MFs were deposited at the subsequent stage of cell wall formation. The first well ordered MFs were oriented in an S-helix. The well ordered MFs had already been deposited at the tracheids where birefringence was first detected under the polarizing light microscope. These results indicate that the deposition of the well ordered MFs, namely, the formation of the secondary wall, begins before the cessation of cell expansion of tracheids. Therefore, it seems that the expansion of tracheids is restricted by the deposition of the secondary wall because the cell walls become rigid simultaneously with the development of the secondary wall and, therefore, the yield point of cell walls exceeds the turgor pressure of the cell. Received: 3 July 1996 / Accepted: 24 September 1996     

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。     

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.     

Coelomomyces psorophorae vartasmaniensis Couch+Laird (1988) (Coelomomycetaeceae: Blastocladiales), a fungal pathogen of the mosquitoAedes australis

The germination of sporangia inCoelomomyces psorophorae vartasmaniensis (C. p. tas.) is uncoordinated and thus there is a mixture of developmental stages in any given population. Continuous urografin gradients separated out the critical stages of germinating sporangia giving four bands, each band representing a consecutive stage of germination. These stages were investigated for changes in the sporangial wall using Transmission Electron Microscopy (TEM). The sporangia have a typical two-layered wall, an electron dense outer layer which can be divided into three distinct sub-layers D1, D2, and D3 and an inner electron transparent secondary wall. Stage 3 sporangia have an intact D1 layer on their outer wall. In the subsequent stages (4 & 4b) there is a progressive breakdown of this layer.     

New Discovery of Palaeostachya from Lower Shihhotse Formation of Taiyuan in Shanxi

Ch. E. Weiss established the genus Palaeostachya in 1876, This genus represents the strobilus of Calamites. As a kind of reproductive organ, the Palaeostachya was first proposed to name Volkmannia by Sternberg but later had been renamed to Palaeostachya by Weiss (1876). In this paper two new species of Palaeostachya, Palaeostachya plagiobracteata sp. nov. and Palaeostachya densibrateata sp. nov., were collected from Lower Shihhotse formarion of Taiyuan in Shanxi Province, and they are described here. Palaeostachya plagiobracteata sp. nov. is a very interesting strobilus, of which both basal and upper parts are not preserved. The specimen of the strobilus is about 2.5 cm long and 1.2 cm in the greatest dismeter. Bracts which extend horizontally from the axis and then turn upward are well preserved. Two rows of them are rather conspicuously. Bract is about 4 mm. long, upper part is about 1 mm. Axis is about 2 mm in width. Four rows of sporangia are distinctly around the axis. The sporangia are sessile, rather large, oblong-ovate in shape, and attach directly to the basal part of bract in an axillary position. Horizon and Locality: Lower Shihhotse formation. Tungshan, Taiyuan, Shanxi. Palaeostachya densibracteata sp. nov. is an important strobilus, of which basal and tip parts are very well preserved. The basal part of sterigma of strobilus is preserved. There are many lepidosomes on the sterigma. Lepidosome is about 3 mm in width and its tip is well preserved. Strobilus is similar to ear of corn in shape about 3 cm long, 1.5 cm. in width. Bracts are well preserved. Six rows of them are more conspicuously in the specimen. The bracts extend horizontally from the axis and then turn upward nearly parallel with the axis. The tip part of each bract appears short triangle, but basal part is contiguous. Sporangia are not preserved in the specimen. Horizon and Locality: Lower Shihhotse formation; Tungshan, Taiyuan, Shanxi.     

SULLITHECA DACTYLIFERA GEN. ET SP. N.: A NEW MEDULLOSAN POLLEN ORGAN AND ITS EVOLUTIONARY SIGNIFICANCE

The pteridosperm (Medullosaceae) pollen organ Sullitheca dactylifera gen. et sp. n. is described from middle Pennsylvanian coal balls. The proximally fused units of the obpyriform compound synangium separate and extend distally as finger-like projections. Each projection contains 4–6 vertically oriented cylindrical sporangia arranged in pairs along the radius of the unit; each unit extends from the outer cover wall toward the center. The distal portion of the compound synangium is hollow as a result of the lateral separation of the centripetally and distally directed synangial units. About 40 tubular sporangia are present in all and dehiscence occurs along a lateral slit in each sporangium. Vascular strands are disposed around the periphery of the organ in addition to a single strand paralleling each sporangium. Two- or three-cell trichomes and stomata are present on the organ surface. Pollen of the Monoletes type is present. A paired row of sporangia in Sullitheca composing a synangial unit is considered the homologue of a paired row of sporangia in the more compact and highly evolved genus, Dolerotheca.     

A NEW SPECIES OF ZOSTEROPHYLLUM FROM THE EARLY DEVONIAN OF NEW BRUNSWICK

A new species of Zosterophyllum, Z. divaricatum Gensel, is described from the late Early Devonian (Emsian) of northern New Brunswick, Canada. It is a Platyzosterophyllum type, consisting of slender sometimes bifurcating axes with laterally borne sporangia oriented to one side of the axis. The species is distinctive in that axes bifurcate within fertile regions and in sporangium shape and attachment. Aspects of the morphology of axis and sporangium cuticle, tracheids, and spores are presented and considered in relation to comparable features in other Zosterophyllum species. Associated vegetative axes exhibiting H- and K-branching patterns and also cuticular features similar to the fertile specimens are described and it is suggested that they may represent parts of the same plant. Zosterophyllum divaricatum is most similar to Z. llanoveranum, Z. fertile, and Z. spectabile, and also resembles Rebuchia ovata to some extent. Z. divaricatum offers considerable information on variation within one species concerning sporangium shape, attachment, and distribution and expands the known diversity of Platyzosterophyllum types.     

A new euphyllophyte Kunia venusta gen. et sp. nov. from the Middle Devonian of Yunnan,South China

Abstract Kunia venusta gen. et sp. nov. is reported from the late Middle Devonian (Givetian) Haikou Formation near Kunming City, Yunnan Province, China. This plant has three orders of naked axes that divide pseudomonopodially. The second‐ and third‐order axes occur in a helix. Fertile appendages are distantly spaced and helically inserted to the third‐order axis; they comprise equally dichotomous branches terminated by two clusters of paired and fusiform sporangia. Sterile appendages are dichotomous and distally recurved. A comparison is made with the basal euphyllophytes including the trimerophytes, cladoxylopsids, zygopterids, stauropterids, and some relevant genera of uncertain affinity. The new plant resembles them in dichotomous appendages with terminal elongated sporangia, but differs mainly in the three orders of pseudomonopodial axes bearing helical laterals. It is thus placed in the Euphyllophytina as incertae sedis. It is suggested that an evolutionary divergence in the branching pattern and appendage morphology might have occurred in the Middle Devonian euphyllophytes, that is, maintaining three dimensions versus yielding more or less planation.     

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.     

STRUCTURE AND COMPOSITION OF THE RESISTANT SPORANGIAL WALL IN THE FUNGUS ALLOMYCES

The fine structure and chemical composition of the wall of resistant sporangia of Allomyces neo-moniliformis were investigated. Studies with the electron microscope showed that the wall is approximately 1.3 μ in thickness and is of complex construction. It consists essentially of three parts: a five-layered outer wall, two layers of “cementing substances,” and a single-layered inner wall. The presence of a highly convoluted cell membrane was also demonstrated. Six structural components were found to make up the walls of the resistant sporangia: glucose, glucosamine, chitin, melanin, protein, and lipids. Comparison of the structure and composition of the walls of resistant sporangia with the walls of hyphae and zoospores of Allomyces as reported by other investigators showed that, while the structure is very different, the composition is quite similar with only melanin and lipids apparently being absent from the zoospore and hyphal walls.     

Xylem of early angiosperms: Novel microstructure in stem tracheids of Barclaya (Nymphaeaceae)

Pit membranes of stem tracheids of all recognized species of Barclaya, an Indomalaysian genus of Nymphaeaceae, were studied with scanning electron microscopy (SEM). Pit membranes of the tracheids are composed of two thick layers, both constructed of fibrils much larger than those of tracheary elements of angiosperms other than Nymphaeaceae. The outer (distal) layer, which comprises the continuous primary wall around the tracheids, is spongiform, perforated by porosities of relatively uniform size, and confined to or most prominent on end walls of stem tracheids. The second layer consists of thick widely spaced fibrils that are oriented axially and are laid down proximally (facing the cell lumen) to the first (outer) layer, although continuous with it. These axial fibrils are attached at their ends to the pit cavities. This peculiar microstructure is not known outside Nymphaeaceae except in Brasenia and Cabomba (Cabombaceae, Nymphaeales), and has not been previously described for Barclaya. The longitudinally oriented threads and strands in perforation plates of secondary xylem of wood and stems of a variety of primitive woody angiosperms (e.g., Illicium) are not homologous to the pit membrane structure observed in stem tracheids of Barclaya, which, like other Nymphaeaceae, has only primary xylem and no perforation plates. The tracheid microstructure reported here is different from pit structures observed in any other group of vascular plants, living or fossil. The tracheid stem microstructures of Barclaya and other Nymphaeaceae appear to be a synapomorphy of Nymphaeaceae and Cabombaceae, and need further study with respect to ultrastructure and function.     

STRUCTURE OF TRACHEIDS IN THREE SPECIES OF LYCOPODIUM

The structure of tracheids in Lycopodium lucidulum, L. clavatum, and L. tristachyum was studied with the light microscope. Protoxylem development is at least sometimes and possibly always mesarch in indeterminate axes of all three species. Centrifugally formed protoxylem elements are reticulate and discontinuities in the secondary walls of these elements are sometimes conspicuously bordered. Wall thickenings of first formed protoxylem elements consist mainly of indirectly connected rings. Late centripetally formed protoxylem elements and transitional elements have a reticulate secondary wall pattern. The narrowest metaxylem elements have circular bordered pits while in wider metaxylem elements pits are bordered and may vary from circular to scalariform. Pitting is uniseriate to triseriate in tracheids of all three species, and intermittent tetraseriate pitting was occasionally observed in L. lucidulum. Crassulae occur in tracheids of the three species, and in L. clavatum an additional framework, probably representing thickened compound middle lamella, is also present. Pits often appear helically arranged, and in all three species pits are connected by thin areas in the secondary wall. Macrofibrils approximately 0.5 μ wide were observed in tracheids of the three species. In L. clavatum the arrangement of macro-fibrils was predominantly bidirectional.     

VEGETATIVE AND FERTILE STRUCTURES OF CYATHOTHECA VENTILARIA FROM THE UPPER PENNSYLVANIAN OF THE APPALACHIAN BASIN

Permineralized stems, leaves and a fertile structure assignable to Cyathotheca Taylor are described from the Late Pennsylvanian Duquesne Coal of eastern Ohio. The new material, C. ventilaria sp. nov., provides the first evidence of vegetative structures for the genus. Vegetative parts are referred to the fertile fragments based upon distinctive vascular morphology, common histological features, and close association. Stems are up to 5.0 mm in diam and have an apparently endarch dictyostele with scalariform metaxylem tracheids. Secondary xylem consists of tracheids with bordered pitting and uniseriate, parenchymatous rays 1–9 cells high. Leaves are arranged in a 3/8 pattern. They are small, pinnately-lobed, and vascularized by an U-shaped bundle. Distal to divergence, one of the apparent leaf traces becomes radial to resemble the base of a fertile structure. This implies a mode of attachment for the fructification and suggests evidence to interpret its homologies. The fertile specimen consists of laminae that diverge from a short stalk, and bifurcate distally. Laminae bear sporangia 0.6–0.7 mm in diam attached adaxially by vascularized pedicels. Spores average 36 μm in diam and are of the Kewaneesporites type. The combination of features now known for Cyathotheca exclude it from assignment to a currently-recognized major group of vascular plants, thus emphasizing that the Pennsylvanian coal swamp flora included a greater diversity than commonly is interpreted.     

MORE EVIDENCE FOR CONIFER DIVERSITY IN THE UPPER TRIASSIC OF NORTH CAROLINA

An ovulate strobilus from the Upper Triassic Deep River Basin, North Carolina, has helically arranged, loosely aggregated, elongated, spatulate bracts with axillary ovule-bearing appendages with about 8–10 ovules attached in two lateral rows, with outwardly directed micropyles. The axillary ovuliferous appendage is homologous with the voltzialean fertile dwarf shoot, but probably not directly evolved from it. More credible is a suggested origin from a completely fertile axillary appendage such as that of the Lower Permian Trichopitys. The occurrence of this cone, Metridiostrobus palissyaeoides, gen. and sp. nov., along with Compsostrobus neotericus and Voltzia andrewsii, reflects considerable diversity among conifer ovulate cones during the Upper Triassic.