A new Neocalamites (Sphenophyta) with prickles and attached cones from the Upper Triassic of China

Remains of the extinct sphenophyte (horsetail) Neocalamites are most widespread in the Middle–Upper Triassic and are typically represented by stem and leaf fragments. Here we report on spectacular new finds of Neocalamites from the Late Triassic Yangcaogou Formation in Liaoning Province, China that include bedding surfaces dominated by nearly complete aerial stems with attached leaf whorls and rare bractless cones. They reveal a monopodial growth habit for the stems, which are covered with downward projecting prickles that probably provided protection against herbivores. These features provide the basis for a new proposed species, Neocalamites horridus. The nodes bear whorls of very long leaves mainly free to their bases, and one specimen bears an attached cone on a long peduncle. Identical dispersed cones have also been recovered. The leaves of adjacent monopodial stems most likely interlocked to support growth in large stands akin to the role now played by branches in large modern Equisetum species. The new Chinese Neocalamites is among the most confidently reconstructed species, and indicates a greater diversity of sphenophyte morphology during the Mesozoic than previously realized.     

OCCURRENCE OF INTERCALARY AND UNINTERRUPTED MERISTEMS IN EQUISETUM

Quantitative data on cell length and mitotic index in elongating internodes of seven species of Equisetum are presented as evidence for the occurrence of two patterns of internode development in rhizomes of different species. In rhizomes of three species of subgenus Equisetum (E. arvense, E. diffusum, E. telmateia) uninterrupted meristems are present, characterized by acropetal internode maturation. In rhizomes of four species of subgenus Hippochaetae (E. hyemale, E. variegatum, E. scripoides, and E. laevigatum) intercalary meristems are present. The number and locations of intercalary meristems are described for an aerial shoot of E. diffusum. The absence or diminution of intercalary meristems from rhizomes of a variety of vascular plants with intercalary meristems in aerial shoots is discussed from the standpoint of adaptive significance.     

EVOLUTION OF COTYLEDONARY AND NODAL VASCULATURE IN THE JUGLANDACEAE

Cotyledonary nodal patterns of the Juglandaceae range from 1-gap, 2-trace to multi-gap, multi-trace. The development of increased nodal complexity is associated with at least two independent evolutionary shifts from epigeal to hypogeal germination. The taxa with epigeal germination such as Engelhardia sect. Engelhardia, Engelhardia sect. Psilocarpeae, Platycarya, and Pterocarya all have 1-gap, 2-trace nodes. The change to hypogeal germination in Engelhardia sect. Oreomunnea and Alfaroa is correlated with the development of 1-gap, 3-trace cotyledonary nodes. The second line has led to large, heavy-fruited members with hypogeal germination and complex cotyledonary nodes ranging from 2–6 gaps. The diversity of nodal patterns is the result of variation on a common theme; five basic vascular strands in the cotyledon, undergoing variations in dichotomy, fusion, and separation, are associated with one to many gaps. Presumably the complex development of the cotyledonary node is a response to increased functional demands of hypogeous cotyledons.     

A FOSSIL EQUISETUM SP. (FAMILY EQUISETACEAE,SUBGENUS HIPPOCHAETAE) FROM THE LATE TERTIARY ASH HOLLOW FORMATION OF NEBRASKA

Silicified leaf-sheath fragments of Equisetum sp. are described from Late Tertiary strata in west-central Nebraska. The close affinity of these fossils to the subgenus Hippochaetae is supported by the presence of sunken stomata arranged in two regular rows on each leaf-sheath segment. This is the first report of the genus Equisetum from the late Tertiary Ash Hollow Formation of central North America.     

An extensin-rich matrix lines the carinal canals in Equisetum ramosissimum, which may function as water-conducting channels

Background and Aims

The anatomy of Equisetum stems is characterized by the occurrence of vallecular and carinal canals. Previous studies on the carinal canals in several Equisetum species suggest that they convey water from one node to another.

Methods

Cell wall composition and ultrastructure have been studied using immunocytochemistry and electron microscopy, respectively. Serial sectioning and X-ray computed tomography were employed to examine the internode–node–internode transition of Equisetum ramosissimum.

Key Results

The distribution of the LM1 and JIM20 extensin epitopes is restricted to the lining of carinal canals. The monoclonal antibodies JIM5 and LM19 directed against homogalacturonan with a low degree of methyl esterification and the CBM3a probe recognizing crystalline cellulose also bound to this lining. The xyloglucan epitopes recognized by LM15 and CCRC-M1 were only detected in this lining after pectate lyase treatment. The carinal canals, connecting consecutive rings of nodal xylem, are formed by the disruption and dissolution of protoxylem elements during elongation of the internodes. Their inner surface appears smooth compared with that of vallecular canals.

Conclusions

The carinal canals in E. ramosissimum have a distinctive lining containing pectic homogalacturonan, cellulose, xyloglucan and extensin. These canals might function as water-conducting channels which would be especially important during the elongation of the internodes when protoxylem is disrupted and the metaxylem is not yet differentiated. How the molecularly distinct lining relates to the proposed water-conducting function of the carinal canals requires further study. Efforts to elucidate the spatial and temporal distribution of cell wall polymers in a taxonomically broad range of plants will probably provide more insight into the structural–functional relationships of individual cell wall components or of specific configurations of cell wall polymers.     

ANATOMY OF NODES VS. INTERNODES IN COLEUS: THE NODAL CAMBIUM

Secondary growth begins in the nodal regions before the internodal regions in Coleus, so that longitudinally discontinuous vascular cambia are formed in the 6th through the 9th or 10th nodes, where the internodal cambium becomes continuous between nodal cambia. The nodal cambia are identifiable by radial seriation in interfascicular regions, typical cytology of fusiform initials, and the presence of a ray system. Anatomical features distinct from the primary plant body are shared by the nodal and internodal cambia. Branching of primary vascular strands, restricted to procambium and phloem, is virtually confined to nodal regions. In secondary growth, vascular branching of xylem and phloem occurs in both nodes and internodes. Xylem strand branches are formed only from derivatives of vascular cambia. It is proposed that the cambium provides the secondary plant body an efficient channel for lateral auxin transport, by which branching across interfascicular regions is facilitated.     

COMPARATIVE MORPHOLOGY OF THE COCHLOSPERMACEAE. II. ANATOMY OF THE YOUNG VEGETATIVE SHOOT

The generic scope and systematic position of the Cochlospermaceae were evaluated using observations from the anatomy of the stem, node, and leaf. There are few basic differences in vegetative anatomy between Amoreuxia and Cochlospermum. Secretory cells and canals, dilated phloem rays, and banded phloem are unifying features. Mature nodal anatomy is 3-trace, trilacunar, and the leaves of both genera have elongate, unicellular, branched idioblasts in the spongy mesophyll. Bixa has some features in common with Amoreuxia and Cochlospermum but is distinctive in vascularization of the petiole, leaf anatomy, and vestiture. Rhopalocarpus is quite different from the above genera, and its placement in a separate family is justified on anatomical grounds. The Cochlospermaceae, consisting of Amoreuxia and Cochlospermum, seem more closely related to the Sterculiaceae and Tiliaceae than to the Flacourtiaceae, Cistaceae, or Violaceae.     

A NEW SPECIES OF PICEA BASED ON SILICIFIED SEED CONES FROM THE OLIGOCENE OF WASHINGTON

Picea eichhornii n. sp. is described from anatomically preserved seed cones. The fossils are from the Early Oligocene Jansen Creek Member of the Makah Fm. which is exposed along the northern shore of the Olympic Peninsula, Washington. The cones are at least 5.5 cm long and up to 3.5 cm in diameter. The cone axis is 4–6 mm in diameter and contains a pith made up of thick-walled parenchyma cells. Resin canals occur in a single ring in the secondary xylem in some specimens but are absent in others. The cortex is mostly parenchymatous and contains numerous large axial resin canals that branch to supply the bract and scale. Vascular traces to each scale and its subtending bract diverge separately from the vascular cylinder of the cone axis. The bract is tongue-shaped and keeled at its base. It is 5 mm wide and up to 9 mm long. The bract trace fades out before entering the bract base while two resin canals extend into the bract base. The ovuliferous scale is about 2.3 cm long and has a thin, probably papery, apex. Resin canals of the scale occur abaxial to the vascular tissue in the scale base, but some bend around the margins of the vascular strand to become adaxial outward. About 20 resin canals occur in the abaxial scale sclerenchyma, and this is the main anatomical feature that distinguishes these cones as a new species. There are less than 14 such canals in cones in a reference collection of 15 modern species and in the two fossil species known from anatomically preserved material. While the new species adds to our knowledge of the diversity of Cenozoic Picea, its affinities within the genus remain undetermined.     

THE ROOT APICAL MERISTEM OF EQUISETUM DIFFUSUM: STRUCTURE AND DEVELOPMENT

The root apical meristem of Equisetum diffusum Don has a prominent four-sided pyramidal apical cell with its base (distal face) in contact with the root cap. Derivatives (merophytes) that contribute to the main body of the root are produced from the three proximal faces of the apical cell. The first division of a proximal merophyte is periclinal to the root surface separating a small inner cell from a larger outer cell. The inner cell is the precursor of the vascular cylinder. The larger outer cell is the precursor of the epidermis, cortex, endodermis, and pericycle. Radial sectors, established early in the development of the cortex, alternate with sectors in the vascular cylinder. These developmental steps show quite clearly that early root development in Equisetum is markedly different from that of most ferns.     

THE ANATOMY AND THREE-DIMENSIONAL MORPHOLOGY OF ANNULARIA HOSKINSII SP. N.

Foliage attached to calamitean shoots is described from coal ball petrifactions of Middle and Late Pennsylvanian age. Leaves correspond to the compression-impression genus Annularia and thus represent the first attached members of this genus to be recognized as petrifactions. Individual leaves contain a single unbranched vascular bundle flanked by wide lateral laminar areas which occupy more than half the leaf cross sectional area. Stomata are confined to broad bands within concave furrows which parallel the vascular bundle on the abaxial leaf surface. Epidermal cells within these furrows are in rows aligned obliquely to the leaf axis, and the rows angle outward at a slight angle towards the leaf margin. Convolutions of the leaf-bearing axes result from nodal diaphragms which are oblique to the shoot axis. Whorled leaves apparently radiate outward in the plane of each obliquely positioned nodal diaphragm. This petrified material helps explain the apparent flattening of entire nodal diaphragms and leaf whorls within the same plane as seen in compression specimens. Annularia hoskinsii sp. n. is proposed, and the systematic position of structurally preserved Annularia foliage relative to the genus Dicalamophyllum is discussed.     

COMPARATIVE STUDIES OF THE NODAL AND VASCULAR ANATOMY IN THE NEOTROPICAL CYATHEACEAE. I. METAXYA AND LOPHOSORIA

Comparative studies of the nodal and vascular anatomy in the monotypic genera Metaxya and Lophosoria are discussed as they relate to the taxonomy and phylogeny of the Cyatheaceae. Both genera are distinctive and primitive with respect to habit, stem and petiole indument, stelar pattern, and nodal anatomy. Metaxya possesses a prostrate, dorsiventral rhizome, whereas a short, upright radial stem occurs in Lophosoria. Trichomes occur on the stems and leaf petioles of these genera. Both Metaxya and Lophosoria have a spiral phyllotaxy, and adventitious buds occur on the petiole bases. The stelar pattern is basically a siphonostele, although frequently a dictyostele is found in Lophosoria. Accessory bundles are lacking in both genera. A characteristic petiole pattern is found in these genera, with an increase in complexity from an undivided strand in Metaxya to the three-parted petiole pattern in Lophosoria. Data from nodal and vascular anatomy indicate that these taxa are distinct from the other genera in the Cyatheaceae and belong in an independent position at the base of the Cyatheoid line, although in some respects an affinity to members of the Dicksoniaceae is indicated.     

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.     

THE VASCULAR TRANSITION REGION OF HELIANTHUS ANNUUS (COMPOSITAE). II. EFFECT OF IONIZING RADIATION

The vascular pattern in the transition region of Helianthus annuus L. may be altered by ionizing radiation in dosages ranging from 650–19,200 R. The alteration of the vascular cylinder is expressed by (1) the failure of normal tracheary elements to mature and (2) by the induction of accessory xylem and/or phloem strands. The anatomical alterations in the vasculature of the transition region are presented and their significance discussed in terms of seedling survival and the mechanisms controlling vascular differentiation.     

Non invasive imaging of water flow in plants by NMR microscopy

Summary Water flow in the stems of the horsetailEquisetum hyemale, the flowering plantStachys sylvatica and the mossDendroligotrichum dendroides was observed non-invasively using NMR microscopy. A Pulsed Gradient Spin Echo sequence using a single phase encoding step was used. We demonstrate by this method that water flow inEquisetum occurs in the carinal canals and in the xylem vessels of the vascular tissue ofStachys and in the central water-onducting strand ofDendroligotrichum.Abbreviations CPU central processing unit - ID internal diameter - NMR nuclear magnetic resonance - RF radio frequency - TS transverse section     

Reproductive biology and population dynamics of wild gametophytes ofEquisetum

This is d study of the reproductive biology of wild gametophytes of Equisetum arvense L., E. fluviatile L., and E. palustre L., which form extensive populations during the summer months on previously inundated bare mud along the margins of reservoirs and lakes in Northern England and Wales. The garnetophytes have all the characteristics of pioneer species: they rapidly attain sexual maturity and are adversely affected by competition. Early in the summer mixtures of male and female gametophytes are produced. The former are smaller and their frequency (which is always less) diminishes throughout the season as a result of bryophytic competition. The rarity of bisexual, initially female, gametophytes, in wild populations is related to the absence of metabolite accumulation which mediates this change in rulture. Significantly different sex ratios between populations and species underlines the labile nature of the sex-determining mechanism in Equisetum. The high frequency of females bearing sporophytes indicates that intergarnetophytic fertilization is highly effective in nature. Absence of any correlation between the incidence of fertilized females and the proportions of males, in conjunction with a consideration of the male gamete dispersal distance, suggests that sporophyte formation is restricted by the availability of ripe archegonia. Sexual reproduction in Equisetum is probably limited by the narrow range of conditions under which gametophytes can become established rather than availability of water for fertilization. The majority of females, which bear one sporophyte, are smaller than unfertilized or polyembryonic females. Their small size results from nutrient demands and allelopathic compounds from the sporophytes which also probably prevent the establishment of gametophytes within mature stands of the parent species. Correlations between female diameters and frequencies of males suggest that gametophytes are more likely to produce archegonia under favourable conditions. The natural reproductive biology of Equisetum is in accord with predictions based on an understanding of the mechanism of sex determination in axenic culture. Several striking parallels between sexuality in Equisetum and dioecious angiosperms are revealed. The absence of winter flooding at two of the gametophyte sites led to the establishment of mature rtands of Equisetum, which produced cones after 3–4 years; two hybrids, E. fluviatile×E. arvense and E. fluviatile×E. palustre, were detected.     

COMPARATIVE STUDIES OF THE NODAL AND VASCULAR ANATOMY IN THE NEOTROPICAL CYATHEACEAE. III. NODAL AND PETIOLE PATTERNS; SUMMARY AND CONCLUSIONS

Comparative studies of the nodal and vascular anatomy in the Cyatheaceae are discussed as they relate to the taxonomy and phylogeny of the family. There is in the Cyatheaceae (excluding Metaxya and Lophosoria) a basic nodal pattern consisting of four major phases of leaf trace separations. Abaxial traces arise from the leaf gap margins, and the last abaxial traces from each side of the gap are larger and undergo numerous divisions. Distally adaxial traces separate from the gap margins, and the last adaxial traces are usually larger and undergo multiple divisions. In addition, medullary bundles frequently become petiole strands of the adaxial arc in the petiole. Rarely, cortical bundles form petiole strands in the abaxial arc in the petiole. Leaf gaps of the squamate genera of the Cyatheaceae are fusiform and possess prominent lateral constrictions which result from medullary bundle fusions and the separation of leaf traces. A characteristic petiole pattern is found in all members of the Cyatheaceae. There is an increase in the complexity of the petiole vascular tissue which results in a gradation from the undivided strand in Metaxya, to the three-parted petiole pattern in Lophosoria, and finally to the much-dissected petiole vascular tissue in the advanced genera. Nodal and vascular anatomy data basically support Tryon's phyletic scheme for the family. The Sphaeropteris-Alsophila-Nephelea line shows certain tendencies toward increased complexity of nodal and vascular anatomy, whereas the Trichipteris-Cyathea-Cnemidaria line shows the same anatomical and morphological characters in a direction of increased simplification or reduction.     

VASCULAR ORGANIZATION IN SHOOTS OF CACTACEAE. I. DEVELOPMENT AND MORPHOLOGY OF PRIMARY VASCULATURE IN PERESKIOIDEAE AND OPUNTIOIDEAE

Primary shoot vasculature has been studied for 31 species of Pereskioideae and Opuntioideae from serial transections and stained, decorticated shoot tips. The eustele of all species is interpreted as consisting of sympodia, one for each orthostichy. A sympodium is composed of a vertically continuous axial bundle from which arise leaf- and areole-trace bundles and, in many species, accessory bundles and bridges between axial bundles. Provascular strands for leaf traces and axial bundles are initiated acropetally and continuously within the residual meristem, but differentiation of procambium for areole traces and bridges is delayed until primordia form on axillary buds. The differentiation patterns of primary phloem and xylem are those typically found in other dicotyledons. In all species vascular supply for a leaf is principally derived from only one procambial bundle that arises from axial bundles, whereas traces from two axial bundles supply the axillary bud. Two structural patterns of primary vasculature are found in the species examined. In four species of Pereskia that possess the least specialized wood in the stem, primary vascular systems are open, and leaf traces are mostly multipartite, arising from one axial bundle. In other Pereskioideae and Opuntioideae the vascular systems are closed through a bridge at each node that arises near the base of each leaf, and leaf traces are generally bipartite or single. Vascular systems in Pereskiopsis are relatively simple as compared to the complex vasculature of Opuntia, in which a vascular network is formed at each node by fusion of two sympodia and a leaf trace with areole traces and numerous accessory bundles. Variations in nodal structure correlate well with differences in external shoot morphology. Previous reports that cacti have typical 2-trace, unilacunar nodal structure are probably incorrect. Pereskioideae and Opuntioideae have no additional medullary or cortical systems.     

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.     

The occurrence of a multilayered structure in the sperm of a pteridophyte

Summary A multilayered structure, previously recorded only in bryophytes, is reported in spermatids of three species of Equisetum. It is interpreted as comprising four layers, recalling the Vierergruppe of Marchantia spermatids. Unlike the multilayered structure of bryophyte spermatids, a rather compact organelle, that of Equisetum forms a thin strip extending almost the whole diameter of the cell. As in bryophytes the upper layer of the Equisetum multilayered structure is composed of parallel microtubules, extending far beyond the underlying layers. The microtubular band is considered equivalent to the similar structure seen in a variety of plant spermatozoids, but in Equisetum it is much longer and composed of many more microtubules than reported from any other plant spermatozoid. The morphology of the multilayered structure is related to the large size and short, broad form of Equisetum spermatozoids.