ULTRASTRUCTURE OF DISPERSED AND IN SITU SPECIMENS OF THE DEVONIAN SPORE RHABDOSPORITES LANGII: EVIDENCE FOR THE EVOLUTIONARY RELATIONSHIPS OF PROGYMNOSPERMS

Abstract: The spore Rhabdosporites (Triletes) langii (Eisenack) Richardson, 1960 is abundant and well preserved in Middle Devonian (Eifelian) ‘Middle Old Red Sandstone’ deposits from the Orcadian Basin, Scotland. Here it occurs as dispersed individual spores and in situ in isolated sporangia. This paper reports on a detailed light microscope (LM), scanning electron microscope (SEM) and transmission electron microscope (TEM) analysis of both dispersed and in situ spores. The dispersed spores are pseudosaccate with a thick walled inner body enclosed within an outer layer that was originally attached only over the proximal face. The inner body has lamellate/laminate ultrastructure consisting of fine lamellae that are continuous around the spore and parallel stacked. Towards the outer part of the inner body these group to form thicker laminate structures that are also continuous and parallel stacked. The outer layer has spongy ultrastructure. In situ spores preserved in the isolated sporangia are identical to the dispersed forms in terms of morphology, gross structure and wall ultrastructure. The sporangium wall is two‐layered. A thick coalified outer layer is cellular and represents the main sporangium wall. This layer is readily lost if oxidation is applied during processing. A thin inner layer is interpreted as a peritapetal membrane. This layer survives oxidation as a tightly adherent membranous covering of the spore mass. Ultrastructurally it consists of three layers, with the innermost layer composed of material similar to that comprising the outer layer of the spores. Based on the new LM, SEM and TEM information, consideration is given to spore wall formation. The inner body of the spores is interpreted as developing by centripetal accumulation of lamellae at the plasma membrane. The outer layer is interpreted as forming by accretion of sporopollenin units derived from a tapetum. The inner layer of the sporangium wall is considered to represent a peritapetal membrane formed from the remnants of this tapetum. The spore R. langii derives from aneurophytalean progymnosperms. In light of the new evidence on spore/sporangium characters, and hypotheses of spore wall development based on interpretation of these, the evolutionary relationships of the progymnosperms are considered in terms of their origins and relationship to the seed plants. It is concluded that there is a smooth evolutionary transition between Apiculiretusispora‐type spores of certain basal euphyllophytes, Rhabdosporites‐type spores of aneurophytalean progymnosperms and Geminospora‐/Contagisporites‐type spores of heterosporous archaeopteridalean progymnosperms. Prepollen of basal seed plants (hydrasperman, medullosan and callistophytalean pteridosperms) are easily derived from the spores of either homosporous or heterosporous progymnosperms. The proposed evolutionary transition was sequential with increasing complexity of the spore/pollen wall probably reflecting increasing sophistication of reproductive strategy. The pollen wall of crown group seed plants appears to incorporate a completely new developmental mechanism: tectum and infratectum initiation within a glycocalyx‐like Microspore Surface Coat. It is unclear when this feature evolved, but it appears likely that it was not present in the most basal stem group seed plants.     

A new species of Tetraxylopteris (Aneurophytales) from the Devonian of Venezuela

Recent investigations of the Middle and Upper Devonian Campo Chico Formation, north-west Venezuela, led to the discovery of a new aneurophytalean progymnosperm, Tetraxylopteris reposana sp. nov. The fossils allow the most detailed morphological outline of a species of Tetraxylopteris Beck to date, and indicate there was more variation within the genus than yet recognized. Tetraxylopteris reposana was probably a bushy plant which formed dense interlocking thickets. Its morphology highlights the similarity between Tetraxylopteris and Proteokalon Scheckler & Banks, and the potential synonymy of these taxa is discussed . Furthermore, there is a morphological continuum between the fertile organs of Tetraxylopteris and Rellimia Leclercq & Bonamo. As the first definitive documentation of Tetraxylopteris in Gondwana, this study indicates aneurophytalean progymnosperms were more widespread than formerly understood. The diversity of the Campo Chico Formation floras and similarity of their composition to northern palaeocontinent assemblages (e.g. the Catskills Clastic Wedge) support the concept of a largely uniform worldwide vegetation.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 275–303.     

The Early Carboniferous progymnosperm Protopitys: new data on vegetative and fertile structures,and on its geographic and stratigraphic distribution

We review progress made during the last 25 years in our understanding of the Protopityales, Early Carboniferous plants belonging to the extinct group of the progymnosperms. Recent studies support previous observations that the only genus of this order, Protopitys, included large arborescent plant with trunks up to 1 m in diameter. All branch orders had an oval eustele, secondary xylem with small rays and tracheid pitting ranging from circular bordered to scalariform bordered, and vascular traces to lateral appendages emitted in an alternate to subopposite distichous arrangement. Leaf morphology remains unknown. New material also confirms that fertile organs of Protopitys consist of branching systems bearing elongated sporangia terminally. Spores have a perispore and range in two size groups, which has been interpreted as a primitive stage of heterospory. The dense wood and fertile parts of Protopitys are comparable to those of the aneurophytalean progymnosperms of the Devonian, but Protopitys is distinct by its eustelic primary vascular system, and its affinities are still uncertain. The genus is now documented from at least nine localities in Europe, North America and Australia. Recent discoveries also indicate that it was present through the whole Mississippian, from the middle Tournaisian to the Serpukhovian.     

Growth, development, and systematics of ferns: Does Botrychium s.l. (Ophioglossales) really produce secondary xylem?1

Developmental morphology and anatomy of Botrychium s.l. were studied to clarify rhizome ontogeny and patterns of tissue maturation that can be used to test the hypothesis that ferns of the Ophioglossales may represent living progymnosperms. Serial anatomical sections of the rhizomes of B. virginianum and B. dissectum reveal that apical meristematic activity and vascular tissue maturation occur over an extended period of several years and then stop. Most of the xylem consists of radial rows of tracheids and interspersed ray-like xylem parenchyma cells that are similar in these respects to secondary xylem, but pits occur on all tracheid walls as is characteristic of primary xylem. No vascular cambium is initiated in mature primary tissues, nor is there secondary phloem. Radial rows of xylem cells are produced by the direct continuation of divisions that begin at the shoot apical meristem, forming a cylinder of radially aligned procambial cells before the differentiation of protoxylem. Continuing divisions over a period of several years increase the number of thin-walled cells and tracheids in each radial row back to about one internode behind where the current year's frond trace diverges from the rhizome stele. At more proximal levels of the rhizome, procambial cell divisions cease and there is no additional tracheid differentiation. These data reveal that the rhizome matures over an exceedingly long period of several years, but that growth is ultimately determinate, thus supporting hypotheses that the Ophioglossales is more closely related to other groups of living ferns than to progymnosperms and seed plants.     

Following basal stem rot in young oil palm plantings

The PCR primer GanET has previously been shown to be suitable for the specific amplification of DNA from Ganoderma boninense. A DNA extraction and PCR method has been developed that allows for the amplification of the G. boninense DNA from environmental samples of oil palm tissue. The GanET primer reaction was used in conjunction with a palm-sampling programme to investigate the possible infection of young palms through cut frond base surfaces. Ganoderma DNA was detected in frond base material at a greater frequency than would be expected by comparison with current infection levels. Comparisons are made between the height of the frond base infected, the number of frond bases infected, and subsequent development of basal stem rot. The preliminary results suggest that the development of basal stem rot may be more likely to occur when young lower frond bases are infected.     

Paleozoic seed fern pollen organs

Paleozoic pollen organs exhibit numerous morphological forms that have been arranged in categories based on their probable organization. Progymnosperm ancestors are characterized by three dimensional branching systems bearing pairs of terminal sporangia. Early Mississippian examples of seed fern fertile branches appear little modified from the progymnosperms. These pteridosperm microsporangia are nonsynangiate and thin walled with longitudinal dehiscence. By Upper Mississippian time all forms show sporangial clustering into large or small groups, with several taxa exhibiting radially symmetrical synangia. In the Lower Pennsylvanian all pollen organs are synangiate and appear to consist of a uniseriate ring of sporangia that either surround a central hollow, or are bilaterally flattened. Sporangial dehiscence in all forms is longitudinal and toward the center of the synangium. In bilateral synangia with no central hollow, the sporangia either separate laterally or effective dehiscence areas are restricted to the free apical portions of the sporangia. Callistophytacean synangia resemble the lyginopterid type, but are abaxial on laminar foliage. This family is thought to have evolved from the lyginopterids during the Early Pennsylvanian. Middle Pennsylvanian medullosan pollen organs are all radial and may be solitary, aggregated into groups, or fused into a large compound synangium. Several pollen organ types are reinterpreted, and the possible evolutionary relationships among the various Paleozoic pollen organ forms are discussed based on synangial organization, patterns of frond branching, and pollen or prepollen morphology.     

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.     

Allometric estimation of total leaf area in the neotropical palm Euterpe oleracea at La Selva, Costa Rica

We estimated the magnitude of the total leaf area of the neotropical palm Euterpe oleracea and examined its allometry relative to the variation in stem height and diameter at La Selva Biological Station in Costa Rica. The allometric relationships between frond leaf area and frond length (from tip to base), and between frond leaf area and number of leaflets, were determined by natural logarithmic regressions to estimate the total area of each frond. Palm total leaf area was then estimated by adding the area of the composing fronds. We fit 14 separate regression models that related one or more of the morphological variables (number of fronds, diameter at breast height, stem height) to the total leaf area. Our results show that palm total leaf area is directly proportional to the number of fronds and palm size as reflected in stem height and diameter. Eight out of the 14 models had r ² values of >0.90 and incorporated a diverse combination of predictor variables. Simple linear regression models were more congruent with the observed values of total leaf area, whereas natural logarithmic models overestimated the value of total leaf area for large palms. Both approaches show a high degree of association among morphological characters in E. oleracea supporting the hypothesis that palms behave like unitary organisms, and are morphologically constrained by the lack of secondary meristems. To afford attaining canopy heights, woody palms need to show a high degree of phenotypic integration, shaping their growth and allometric relationships to match spatial and temporal changes in resources.     

Review of the Fossil Evidence for the Origin and Earliest Evolution of the Seed-plants

The earliest identified seed-plants are found in Latest Devonian aged sediments from Northwest Europe and North America and are believed to have evolved from within the progymnosperms, a group of anatomically advanced pteridophytic plants. However, current evidence makes it problematic to determine from which particular progynmospermous lineage the seed-plants evolved, with the major contender being the Aneurophytalean progymnosperms. The evidence for the ancestral stock to the seed-plants is summarised and the morphologies of the earliest known seed-plants are considered. From their first geological occurrence, the seed-plants are morphologically diverse to such an extent that the identification of a single potential ancestral morphology is impossible. In the light of the evidence so far presented, the precise origin of the seed-plants is unresolved and in need of new evidence relating to the progynmosperm/seed-plant transition. Future lines of research are also suggested.     

种子植物的起源及其早期演化

迄今为止世界上最早的种子植物发现于北美和欧洲西北部的晚泥盆纪地层。这些种子植物可能起源于前裸子植物，即解剖构造上较为进化的一类蕨类植物。然而，从目前的证据来看尚无法确切判断这些种子植物起源于前裸子植物中的某个特定类群，比如古羊齿目和戟枝木目。作者总结了种子植物祖先的有关证据，并对目前已知的最早的种子植物的形态进行了概述。由于种子植物自出现起即表现出丰富的形态多样性，因而尚无法对某个可能的祖先形态作出判断。目前的研究尚不能确切回答种子植物起源的全部问题，仍需寻找有关前裸子植物与种子植物过渡类型的新证据。作者对未来的研究提出了建议     

Phyllotaxis in the Oil Palm: Applications in Selection of Interspecific Hybrids

The main purpose of the investigation was to determine whethermeasures associated with or dependent on the differences inphyllotaxis organization of the stem, fronds, and inflorescencesof three families of oil palms which differ considerably ingrowth habit and genetic background could be associated with,or allow prediction of their yield differences. Two of the familiesused are interspecific crosses between Elaeis guineensis andE. melanococca and these hybrids yield almost double that ofthe third ‘family’ (a group of E. guineensis selections)and have about twice the individual frond areas, inflorescencevolume, and surface area and about half the rate of verticalgrowth; they have, however, a slightly slower rate of frondand inflorescence production. One of the views expressed isthat the phyllotaxis organization which determines the measuredreduction of stem volume in the interspecific crosses is associatedwith the greater development of more productive reproductiveelements. Further, it is suggested that measurements of frond(even in the juvenile stage) and both male and female inflorescenceorganization could be used in yield prediction for these familiesand possibly other genetic groups-and thus increase the speedand efficiency of genetic selection.     

Non-timber forest product harvesting in alien-dominated forests: effects of frond-harvest and rainfall on the demography of two native Hawaiian ferns

Non-timber forest products (NTFP) represent culturally and economically important resources for millions of people worldwide. Although many NTFP are harvested from disturbed habitats and therefore subject to multiple pressures, few quantitative studies have addressed this issue. Similarly few NTFP studies have assessed seasonal variation in demographic rates even though this can confound harvest effects. In Hawaiȁ8i, the wild-gathered ferns, Microlepia strigosa and Sphenomeris chinensis, represent highly important cultural resources but declining populations have led to conservation concerns. Both ferns are harvested from disturbed, alien-dominated forests and contemporary Hawaiian gathering practices often consist of harvest and concurrent weeding of alien invasive species. We assessed the effects of concurrent frond-harvest and alien species weeding on frond structure, density, and rates of production by comparing experimentally harvested vs. control plots, and documented relationships between frond demographic patterns and precipitation. Gathering practices had no impact on frond density of either species or on most other demographic parameters over the short term. Exceptions included a significant decrease in the density of the longest S. chinensis fronds and a significant decrease in M. strigosa frond production when fronds were gathered without alien weeding. However, seasonal and annual changes in frond density and production occurred across all plots of both species and were significantly correlated with precipitation. The relatively low harvest effects for both species are likely due to several factors including short frond longevity and the strict criteria used by gatherers to select harvestable fronds. The potential for sustainable harvest in the context of alien-dominated forests is discussed.     

Developmental anatomy of vascular cambium and periderm ofBotrypus virginianus and its bearing on the systematic position of ophioglossaceae

The developmental anatomy of the vascular cambium and periderm ofBotrypus virginianus was studied, and its bearing on the systematic position of Ophioglossacease is discussed. The cambial zone including cambium is initiated in a procambial ring of the stem before primary vascular tissue is well differentiated. The presumed cambium is composed of fusiform and ray initials. The cambium is extremely unequally bifacial, producing secondary xylem centripetally, and quite a small number of parenchymatous cells but no secondary phloem centrifugally. The cambial activity persists long, although it is very low in the mature part of the stem. It seems that the circumferential increase of the cambium is accommodated by an increase in the number of cambial initials. Secondary xylem is nonstoried and composed of tracheids with circular-bordered pits with evenly thick pit membranes, and uniseriate or partly biseriate radial rays. It makes up the bulk of the stem xylem. Periderm is formed almost entirely around the stem, simultaneous with its increment due to the secondary xylem. The combination of these anatomical features of secondary tissue supports the idea that Ophioglossaceae are living progymnosperms.     

RELLIMIA THOMSONII FROM THE GIVETIAN OF NEW YORK: SECONDARY GROWTH IN THREE ORDERS OF BRANCHING

Pyritized axes of the vegetative branching system of Rellimia thomsonii, from the Givetian of New York were described with emphasis on the presence and position of the secondary tissues in relation to the known axis orders. The morphology and anatomy of four axis orders (N, N + 1, N + 2, N + 3) was described using two well-preserved specimens. One specimen showed the largest known (N) axis order, with spirally attached N + 1 axes. The other specimen, from a more distal portion of the plant, showed the N + 2 axis order with spirally arranged N + 3 axes. The shape of the primary xylem, discernible in three of the axis orders (N + 1, N + 2, N + 3), was a three-lobed protostele with mesarch maturation of the primary xylem. The protostele of the N + 3 axis order, with only primary growth, was identical to that of fertile N + 3 axes. The other axis orders (N + 2, N + 1, N) had wood surrounding the primary xylem. The wood of the N+2 axis was limited, and occurred only in the basal portion of the axis, but that of the N+1 and N axes was extensively developed with two and three growth layers, respectively. This wood was pycnoxylic with narrow rays similar to other aneurophytalean progymnosperms. Surrounding the secondary xylem was secondary phloem indicating the vascular cambium in Rellimia was bifacial. The presence of growth layers was discussed in relation to the climatic interpretation placed on their presence and how it correlates with paleogeographic reconstructions of the Devonian.     

ABSCISSION AND THE AXILLARY FROND IN SPIRODELA OLIGORHIZA (LEMNACEAE)

Two abscission zones are present in fronds of Spirodela oligorhiza. In one zone the abscission layer separates the daughter frond and its connecting stalk from the mother frond. The second abscission layer separates the daughter frond from the connecting stalk. An axillary frond is asymmetrically positioned at the base of each daughter frond where it joins the mother frond. In right-handed physiological clones the axillary frond is to the right of the connecting stalk and in left-handed clones, to the left of the stalk. When a daughter frond abscises it leaves behind its axillary frond in the pocket of the mother frond. Histological features of abscission and treatments that induce abscission in Spirodela oligorhiza are described.     

Clinal Variation of Frond Morphology and its Adaptive Implication in the Fern Ceratopteris thalictroides in Japan

Abstract Frond morphology of the highly polymorphic, annual fern Ceratopteris thalictroides was examined for local populations in south and central Japan. Among the characters examined, frond length and frond dissection were most useful for recognizing several forms. Through the comparison of these characters among populations, a geographically clinal pattern of variation was recognized; plants in higher latitudes have smaller and less dissected fronds. Plants usually grow in rice paddies during a limited period from the harvest time to the beginning of the frost season. Concordance was observed between the degree of frond length and dissection of populations and the length of the viable period in their localities. The variability of cultivated progeny of selected populations from the south and the north suggests that there are at least two ecological lines; one is genetically restricted to large and much dissected fronds and the other to smaller and less dissected fronds. The latter may be an adaptation to the short viable period in rice paddies in northern regions. High phenotypic plasticity enables these lines to adjust well the degree of frond length and dissection to the length of the viable period in each locality.     

Neuropteris attenuata, a narrow-stemmed, leaning or lianescent seed fern from the Upper Pennsylvanian of Lower Saxony, Germany

A stem segment and frond of Neuropteris attenuata Lindley et W. Hutton (Pteridospermopsida, Medullosales) are described from the lower Westphalian D (Upper Pennsylvanian) of the Piesberg quarry near Wallenhorst (north of Osnabrück, Lower Saxony, northwestern Germany). The narrow diameter of the stem (3–4 cm), conspicuously long internodes (>20 cm), and the large size of the frond (90 cm long) strongly suggest that N. attenuata was a non-self-supporting, leaning or lianescent plant.     

Fertile and sterile frond segments of the lyginopterid seed fern Feraxotheca

Fertile and sterile frond segments of the lyginopterid seed fern Feraxotheca culcitaus are described. In both the fertile and sterile specimen three orders of frond axes are borne alternately and in the same plane. Antepenultimate and penultimate frond axes are characterized by a C-shaped vascular bundle and numerous canals containing a golden-colored material in the fossilized condition. Synangia with only four sporangia demonstrate radial symmetry; bilateral symmetry is present in larger synangia. Pinnules possess dichotomous venation, with a single vein entering each lobe of the deeply incised lamina. A single canal occurs below each vein, and vein ends have an amplified vascular bundle. Morphologically Feraxotheca compares most closely with the compression genus Crossotheca. The morphology of the vascular bundle in the largest pinna is similar to a Lower Carboniferous specimen of Rhodea.     

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

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