Twin shoots borne on a Botryopteris frond from the British upper carboniferous

A long rachis of Bottyopteris aff. hirsuta bearing twin opposite lateral shoots is described from a coal ball of Westphalian age collected in Lancashire. The simultaneous emission, in the middle region of a frond, of two short axes each bearing a large number of poorly developed leaves is a feature hitherto unrecorded for the genus. The specimen suggests that this Paleozoic fern consisted of long fronds whose branching pattern showed a certain capacity for morphological variation; it provides yet another example of the surprisingly limited development of shoots borne on Botryopteris fronds.     

THE OCCURRENCE OF “ROOT GAPS” IN THE RHIZOME SOLENOSTELE AND LEAF TRACE OF DENNSTAEDTIA CICUTARIA

The vascular system of the rhizome axes of Dennstaedtia cicutaria consists of a solenostele with an amphicribral vascular bundle centrally located in the pith. Each leaf has a single trace which is an amphicribral vascular bundle. At each node of an axis there is a complex consisting of the main axis, leaf base, and a branch axis attached to the basiscopic margin of the leaf base. Numerous roots are present on the rhizomes and the abaxial side of the leaf bases. Parenchymatous gaps occur in the rhizome solenostele and the leaf trace directly above the departure of some of the root traces. These gaps are termed root gaps. In some instances the root gaps are confluent. However, not all of the root traces have an associated root gap. The leaf trace is inserted laterally on the main and branch axes at the node so that the acroscopic leaf trace margin anastomoses with the main axis of the vascular system and the basiscopic margin with that of the branch axis. Two leaf gaps are associated with each leaf trace, one occurring in the main axis solenostele and the other in the branch axis solenostele. The medullary bundle of each axis anastomoses with each leaf trace at its point of attachment to the rhizome solenostele. Thus, the medullary bundle forms a continuous vascular strand from leaf trace to leaf trace in any given rhizome axis.     

THE MORPHOLOGY AND RELATIONSHIPS OF RHODEA,TELANGIUM, TELANGIOPSIS,AND HETERANGIUM

This study deals with four form or organ genera from the Upper Mississippian (Chester Series) of the Illinois Basin, and provides evidence that they were produced by a single natural genus with gymnospermous affinity. The plant remains—compressions, impressions, petrifactions, and specimens that combine compression or impression with petrifaction—allow examination of both external morphology and internal anatomy. The specimens include foliage corresponding to Rhodea, stems and petioles corresponding to Heterangium, and synangiate fructifications corresponding to either Telangium or Telangiopsis. The stems and foliage are considered parts of the same plant because of the identity of the anatomical and cuticular features of petioles attached to stem and those petioles with attached foliage. The fertile material is regarded as part of the same plant because: (1) The anatomy of axes of the fertile specimens is like that of the sterile specimens. (2) A single specimen may contain both sterile Rhodea-type axes and fertile regions. (3) Axes bearing synangia have the same size and patterns of divisions as the sterile foliage. Features that indicate lyginopterid affinities include: (1) Equal forking of the petiole. (2) Presence of fiber bands in the outer cortex and sclerotic nests in the inner part of the cortex. (3) Crowded circular bordered pits on the lateral walls of the metaxylem tracheids. (4) The presence of a small amount of secondary xylem. A variety of structural details of the stem and petiole suggest the genus Heterangium. The phyletic position of the plant that produced Rhodea, Telangium, Telangiopsis, and Heterangium is reviewed in light of such discoveries as the presence of a planated frond that lacks a lamina and the presence of both monolete and trilete microspores in a single synangium.     

Re-evaluation of Halle’s fertile pteridosperms from the Permian floras of Shanxi Province, China

In 1927, T. G. Halle published an extensive synthesis of the Permian fossil plant assemblages from China’s Shanxi Province that included five genera and species of seed plant that either had ovules in organic attachment or closely and repeatedly associated with foliage. These fertile pteridosperms were the subject of a later publication (1929) that provided additional information on each species, but since that time only Nystroemia pectiniformis Halle has been subjected to detailed investigation based on the original materials. Reinvestigation of the remaining four species has yielded additional examples and new information for each species, and has confirmed the pteridospermalean affinity for three of the four species. Alethopteris norinii Halle shows organic attachment of a pollen organ similar to Wittleseya (Newberry) Halle and is reinterpreted as a microsporangiate medullosan pteridosperm. Emplectopteris triangularis Halle and Sphenopteris tenuis (Schenk) Halle both bear numerous ovules attached directly to the leaf laminae, with Emplectopteris conforming with the gross-organisation of callistophytalean pteridosperms and S. tenuis being interpreted as a derived lagenostomalean pteridosperm. However, discovery of the counterpart to the key specimen of Pecopteris wongii Halle that Halle interpreted as bearing ovules demonstrates this species to be a sterile marattialean frond that agrees with its assignment to Pecopteris. In addition, two further specimens are demonstrated to be fertile organs of previously unknown pteridosperms; Norinosperma shanxiensis gen. et sp. nov. bears cardiocarpalean ovules abaxially on the proximal portions of the frond with ovules positioned directly on the veins of a leaf lamina; Norinotheca shanxiensis gen. et sp. nov. bears large pollen organs abaxially and singly on the vein of the leaf lamina. Collectively Halle’s pteridosperms remain important today as they are some of the few specimens that actually demonstrate organic connection of fertile organs to foliage, helping to characterise the morphology and organisation of this seed plant group. These species also demonstrate the persistence of seed plant families and genera in the Permian of China that became regionally extinct at the end of the Carboniferous in Euramerica, and show that pteridosperms remained an important component of Permian as well as Carboniferous floras globally.     

Pollen Organ Telangiopsis sp. of Late Devonian Seed Plant and Associated Vegetative Frond

Pollen organ Telangiopsis sp., associated with but not attached to vegetative fronds, has been collected from the Upper Devonian (Famennian) Wutong Formation, Dongzhi County, Anhui Province, China. Fertile axes with terminal pollen organs are dichotomous for 2–4 times and may be proximally attached by fragmentary pinnules. Pollen organs are synangiate and borne on the top of a short stalk. Synangia are radial in symmetry and each consists of 4–8 elongate microsporangia fused at base. Microsporangia have a longitudinal dehiscence line and show a tapered apex. The associated stem is spiny and bears a vegetative frond which bifurcates once at the basalmost part. Frond rachises possess one order of pinna arranged alternately. Pinnules are borne alternately, planate, highly dissected, and equally dichotomous for 2–3 times. Comparisons among Late Devonian seed plants recognize several branching patterns in the fertile fronds/axes bearing terminal pollen organs. Telangiopsis sp. reinforces that the Late Devonian pollen organs are synangiate usually with basally fused microsporangia. It is suggested that the evolutionary divergence of radial and bilateral symmetries of pollen organs may have occurred in the Famennian, when the earliest seed plants evolved planate and sometimes laminate pinnules.     

Morphology, anatomy and vasculature of leaves in Pinus (Pinaceae) and its evolutionary meaning

The morphology, anatomy and vasculature of Pinus leaves was studied. The results indicate that the Pinus leaf is always supplied with only one single vascular bundle, which is surrounded by a prominent bundle sheath. In several taxa, especially of subgenus Pinus, the vascular bundle may however be subdivided by longitudinal parenchymatic bands in the middle part of the leaf. As a result, the single bundle gets the appearance of two individual bundles, which are surrounded by a common bundle sheath. The general rule that a bundle sheath does sheath only one single bundle, as in other gymno- and angiospermous seed plants, applies therefore also for Pinus. The morphological and anatomical similarities between cladodes of Sciadopitys (Sciadopityaceae) and Pinus leaves are based on a completely different bauplan. The idea of “hidden cladodes” in some Pinaceae is therefore obsolete.     

SYMMETRY AND DEVELOPMENT OF LIMNOBIUM SPONGIA (HYDROCHARITACEAE)

Limnobium spongia produces upright vegetative axes and prostrate stolons. The upright axes bear new stolons, whereas the stolons bear new upright axes and fertile and sterile branching systems. Upright axes and fertile and sterile branching systems are all interpreted to have sympodial growth. However, it was not determined whether growth of stolons is monopodial or sympodial. Both stolons and upright axes branch in alternate plastochrons, and branching is achieved solely by the bifurcation of apical meristems. Each meristematic bifurcation is interpreted to represent the formation of a precocious lateral bud. The upright axes develop from presumed precocious lateral buds on stolons, whereas such buds on upright axes produce renewal shoots. Limnobium spongia exhibits a marked degree of mirror-image symmetry.     

Roles for structural and temporal shelter-changing by fern-feeding lepidopteran larvae

Summary Larvae of the pyralid moth, Herpetogramma aeglealis, construct feeding shelters upon the Christmas fern (Polystichum acrostichoides). Field and laboratory study involving 532 shelters showed that as the larvae mature, they sequentially inhabit approximately 5 shelters of 3 distinct types, constructed at night on different fronds of the same plant. The bundle shelter, simple and ephemeral, is first to be inhabited and constructed. The fiddlehead shelter which houses slightly older larvae strongly resembles contemporaneously emerging frond fiddleheads. The final shelter form, the globe, is a silk-bound ball of leaflets at the frond tip. An individual larva usually constructs 3 globe shelters on different fronds of the same plant before completing its development. As shelter sites, sterile Polystichum fronds are chosen preferentially over fertile fronds. The bundle and fiddlehead shelter forms, less abundant, appear cryptic to humans and perhaps to other vertebrates. The final globe shelter form is larger and quite conspicuous. However, the persistence of empty globe shelters left on the plant as the larva moves to a new one may serve to make searching for larvae less profitable for potential predators and parasites. We suggest that the energetic costs of constructing and occupying multiple shelters may be offset by circumvention of reduced frond palatability and reduced exposure to predators and parasites.     

Morphogenesis in Selaginella. III. Meristem determination and cell differentiation.

Selaginella willdenovii Baker is a prostrate vascular cryptogam with a dorsiventral stem. At each major branching of the stem tip a dorsal and a ventral angle meristem are formed. The ventral meristem becomes determined as a root and the dorsal meristem as a shoot. Indoleacetic acid (IAA) is transported basipetally in the stem and has been found to be the regulatory agent for meristem determination both in vitro and in vivo.Growth measurements of intact plants indicated that the sequence of development for each stem unit is frond expansion, internodal elongation, ventral meristem growth as a root, and dorsal meristem growth as a shoot. The principal experimental findings of this study are as follows. Triiodobenzoic acid (TIBA), an inhibitor of auxin transport alters the normal pattern of development in intact plants, causing ventral meristems to develop as shoots and dorsal meristems to develop precociously. Dorsal meristems grown in sterile culture on an auxin-free medium develop as shoots, but in the presence of IAA develop as roots. Meristems transferred after excision from auxin-free to plus-auxin medium on successive days showed an increasing tendency to develop as shoots, with more than 50% doing so by day 5. The mitotic index is low at the time of excision of the meristem, rises to a peak on day 5 and then declines.     

The Structure and Development of the Thallus in the British Species of Gelidium and Pterocladia

The apical structure and the development of the thalli of allthe British species of Gelidium and Pterocladia have been investigated;the development of G. pulchellum is described in detail. Eachaxis is terminated by one or more apical cells, which by theirsegmentation form the tissues of the thallus. An axial filamentis distinct for a short distance behind each apical cell, butsecondary pit-connexions develop rapidly so that in sectionthe mature axis has the appearance of a multi-axial structure. Lateral branches of the frond develop by the segmentation ofthe lateral branch apical cells, which are formed by the transformationof superficial cortical cells, either in the meristematic ormature parts of the axes. The extreme variability of externalappearance is due principally to the indeterminate origin ofall lateral branches. The thallus in the British species of Gelidium and Pterocladiaconsists of erect fronds borne on creeping axes. The relativeproportions of the frond and creeping axes in various speciesand their survival through adverse conditions are discussed.     

STUDIES OF PALEOZOIC FERNS: THE FROND OF ANKYROPTERIS GLABRA

Eggert , Donald A. (Southern Illinois U., Carbondale.) Studies of Palerzoic ferns: The frond of Ankyropteris glabra. Amer. Jour. Bot. 50(4): 379–387. Illus. 1963—The major features of the frond of A. glabra are described on the basis of preserved parts found in Middle Pennsylvanian coal ball material from Illinois. The frond is planated and has well-developed foliar laminae. Primary pinnae arise from the petiole in 2 alternating series, and secondary pinnae arise in a similar fashion from the primary pinnae. Foliar laminae occur on the secondary pinnae and have dichotomous venation. The xylem of the petiole has a diupsilon configuration in the lower part of the axis, while higher in the petiole the xylem forms a strand resembling that of the European species A. westfaliensis. The xylem strands of the primary pinnae arise from the adaxial antennae of the petiolar vascular strand as somewhat C-shapcd bodies and develop antennae and become H-shaped at higher levels. A gap occurs in the antenna of the petiole vascular system above the level of departure of the primary pinna trace. Terete vascular strands occur in the secondary pinna axes which arise from the adaxial antennae of the xylem of the primary pinnae. The foliar laminae are relatively thin, have an irregular outline, and their histology is like that found in many living ferns. The frond of A. glabra illustrates that leaf evolution had progressed in at least one species of the coenopterid family Zygopteridaceae to the extent that an essentially 2-dimensional frond of modern aspect, and with well-developed foliar laminae, was present by Middle Pennsylvanian time.     

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.     

The primary vascular system of Callixylon

The primary vascular system of Callixylon (main stem axes of Archaeopteris) is composed of a discontinous cylinder of longitudinal bundles from some of which traces diverge radially in a regular, helical pattern. Tangential divisions and fusions of primary xylem strands occur in approximately equal frequency and, thus, the number of bundles in the system remains constant in the specimen studied. Following divergence, traces increase in size and become ribbed distally. The available evidence suggests that all of the traces observed in this study are branch traces. Branch bases, embedded in secondary xylem, have the anatomical characteristics of penultimate axes of lateral branch systems of Archaeopteris. It is suggested that some regions of main axes of Archaeopteris might have borne lateral branch systems only, lacking entirely simple leaves. Lateral branch systems, therefore, might have been more similar to compound leaves than previously supposed.     

New Middle Jurassic record of fern genus Eboracia (Dicksoniaceae,Filicales) and its spatio-temporal distribution in China

As an important component of the Mesozoic flora, the extinct fern genus Eboracia Thomas (Dicksoniaceae, Filicales) is widely reported in China with diverse fossil records. New material of Eboracia lobifolia, represented by a nearly intactly preserved fossil frond, is described herein from the Middle Jurassic Haifanggou Formation in Beipiao of western Liaoning, Northeast China. The frond is lanceolate in gross outline, at least 38.0 cm long, and can be divided into the basal sterile part, the upper fertile part and a transitional part in between. The new discovery confirms for the first time that the frond of E. lobifolia is hemidimorphic rather than holodimorphic. Many in-situ spores were detached from the sori of the fertile pinnae, which are characterized by rounded-tetrahedral shape, smooth surface, distinct border, and a wide, long triradiate crack almost reaching the equator. Comparing with dispersed spores in the same horizon, these in-situ spores seem to be most similar to Cyathidites minor Couper in morphology. A spatio-temporal analysis of Eboracia in China shows that Eboracia with totally four species occurred in a time interval ranging from the Late Triassic to the Early Cretaceous in China, and mostly flourished in the Middle Jurassic; the genus was widely distributed in both the Northern and Southern Phytofloristic Provinces of China, particularly in southern China during the Late Triassic to Early Jurassic, while more abundant and diverse in northern China during the Middle Jurassic to Early Cretaceous.     

An upright psaroniaceous stump and two surrounding pecopteroids from the early Permian Wuda Tuff Flora

An upright Psaronius stump was discovered in the upper part of Taiyuan Formation (Asselian) of Wuda Coalfield, Inner Mongolia, China. The stump is 47 cm high and truncated at the upper part of the ash layer in which it is preserved. At the height of 38 cm, the stem possesses a stelar configuration comprised of 3~4 cycles in 5 orthostichies. The Wuda Psaronius stump rapidly widens at the base until it attains a stable axial diameter; a root mantle is present in the proximal zone of primary body expansion, but not distal to that zone. This architecture is further demonstrated by a prostrate Psaronius trunk almost 7 m long with a stable stem diameter and two rows of frond scars visible over most of its length. This developmental mode is seen among extant leptosporangiate tree ferns but differs from that previously documented among North American Pennsylvanian marattialean tree ferns whereby the primary body of Psaronius stems increases continually in width towards the apex and is buttressed by a thick root mantle that compensates for the obconical nature of the stem. Two types of pecopteroid fronds associated with the Psaronius stump are identified as Pecopteris cf. polymorpha and Pecopteris orientalis based on their pinnule morphology. The Pecopteris orientalis frond possesses a penultimate rachis with stewartiopteroid anatomy and is more likely to be the leaf of the Psaronius stump based on a recently reported Psaronius specimen with stem and frond in connection from the same locality.     

<Emphasis Type="Italic">Istchenkophyton filiciforme</Emphasis> gen. et sp. nov., a new small vascular plant with thick cuticle from the Devonian of Voronezh Region (European Russia)

Remains of a new small vascular plant, Istchenkophyton filiciforme gen. et sp. nov., are described from the Upper Givetian strata of Pavlovsk vicinity (Voronezh Region). The plant is characterized by a very thick cuticle (up to 300 μm), complex branching pattern, similar to that of a frond rachis, anomocytic stomata, and a well-developed vascular bundle with a central elongated in transversal section protoxylem strand and massive metaxylem composed chiefly of scalariform bordered-pitted tracheids. A reconstruction of the new plant is proposed; its ecology and systematic position are considered. In the light of new data relationships of problematic Devonian plants with thick cuticle are discussed.     

胡杨叶片解剖特征及其可塑性对土壤条件响应

表型可塑性对极端干旱区植物个体适应具有重要的意义。该研究选择内蒙古额济纳胡杨林内不同土壤条件下胡杨个体,对其叶片解剖结构与土壤因子关系进行分析,阐明不同土壤条件下叶片解剖结构及其可塑性规律。结果表明：除主脉木质部与维管束面积比外,异质性的土壤条件下胡杨叶片各解剖结构均存在明显的趋异适应;土壤水、盐条件影响了以上表皮细胞为代表的表皮组织的生长发育,土壤肥力影响了以主脉维管束为代表的输导组织以及栅栏组织的生长;生境环境异质性是胡杨叶片解剖结构可塑性的根本原因,解剖结构可塑性使得胡杨在不同环境、同一环境不同结构形成适应性差异,不同土壤条件下叶片主脉木质部、主脉维管束的可塑性较大,是胡杨适应异质性土壤条件的主要结构。叶片解剖结构特征及其可塑性对胡杨适应干旱环境具有重要生态作用。     

The size of the frond in the genus AlethopterisSternberg(Pteridospermopsida,Carboniferous)

A fine specimen of AlethopterisSternberg collected in the North-Pas-de-Calais coalfield allows an estimate of the general size of the frond to be made.     

Ontogeny of the vascular bundles and contiguous tissues in the maize leaf blade

The patterns of initiation and early development of the minor and major veins in the flat portion of the leaf blade of maize (Zea mays L.) follow similar patterns. The veins and their associated bundle sheath cells commonly arise from cell assemblages derived from a single cell lineage, or longitudinal file of cells, rather than from two “half vein units” derived from different cell lineages. In addition, apparently, none of the vascular cells derived from the procambium is directly related ontogenetically to a bundle sheath cell. In veins derived from larger cell assemblages, the lateral bundle sheath cells are more closely related ontogenetically to the mesophyll cells, which are derived from the ground meristem, than to the vascular cells, which are derived from procambium. The bundle sheath cells, accordingly, are interpreted as being ground meristem in origin.