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.     

Dicroidium diversity in the Upper Triassic of north Victoria Land,East Antarctica

We describe an assemblage of well-preserved, cuticle-bearing plant compressions collected from a single bed in the Upper Triassic of Timber Peak, East Antarctica. The flora is dominated by the seed fern Dicroidium, a morphogenus that is widespread and usually dominant in Middle to Late Triassic plant assemblages throughout Gondwana. Despite its common occurrence, the systematic classification of Dicroidium and its individual species remains problematic. This study demonstrates that a combination of macromorphological and cuticular features enables a suitable discrimination of Dicroidium species. The present Dicroidium assemblage is remarkably diverse and dominated by D. elongatum with subordinate proportions of D. odontopteroides, D. crassinervis, and a new species provisionally named D. sp. A. In addition, D. dubium, D. spinifolium, and D. coriaceum occur sporadically. We reinstitute D. spinifolium and propose an emended diagnosis. Other taxa present include putative bryophyte remains, Lepidopteris langlohensis, and Heidiphyllum elongatum. Another collection of plant fossils from a second plant-bearing horizon at Timber Peak consists of fragmentary remains of Cladophlebis sp. and H. elongatum. Epidermal and cuticular features of the Dicroidium fronds suggest that this flora flourished under favourable climatic conditions, although it was situated at latitudes of about 70° S and must have experienced long annual periods of darkness during the austral winter. The cuticles studied point to a short leaf lifespan and very high photosynthetic performance of Dicroidium fronds. We suggest that this leaf phenology has facilitated the Dicroidium plants to colonize the polar latitudes of the Triassic greenhouse world so successfully.     

Comparative leaf anatomy and morphology of some neotropical Rutaceae: Pilocarpus Vahl and related genera

Previous anatomical studies have been restricted to the foliar aspects of Pilocarpus. However, no anatomical studies analyzing the foliar aspects of Pilocarpus in relation to related genera have been carried out. Therefore, the aim of this study was to identify characters for future taxonomic and phylogenetic studies in Rutaceae, particularly in Pilocarpus, and to discuss the characteristics associated with the simple or compound leaf condition for the group. The petiole and the leaf blade of 14 neotropical Rutaceae species were analyzed, and the following characteristics were observed in all leaves studied: stomata on both surfaces; secretory cavities, including mesophyll type; camptodromous?Cbrochidodromous venation pattern; and free vascular cylinder in the basal region of the petiole. Additional promising characters were identified for future taxonomic and phylogenetic studies in the Rutaceae family, especially for the Pilocarpus genera.     

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.     

Limits of stratigraphic distribution of Glossopteris in India: Discussion

The stratigraphic limits of Glossopteris in India have been reviewed by Sarbadhikari (1974) and are inferred to range from the Sakmarian-Artinskian (early Permian) to the Permo-Triassic transition. The lower limit in peninsular India is revised as Asselian with the help of Eurydesma-dominated fauna. Glossopteris was well established during the Asselian in the Salt Range but appeared during the late Sakmarian-Artinskian in the Himalayas. In peninsular India, Glossopteris flourished well during the middle Triassic and continued at least upto the Rhaetic, in association with Dicroidium. The Glossopteris-Dicroidium assemblage thus is not a time-restricted transitional zone. Dicroidium with associated forms characterises a flora which is distinct from the older Glossopteris flora and the younger Ptillophylum flora. A three-fold biostratigraphic subdivision of the Indian Gondwana thus has been advocated.     

MORPHOLOGY AND DICHOTOMOUS VASCULATURE OF THE LEAF OF KINGDONIA UNIFLORA

Foster , Adriance S. (U. California, Berkeley), and Howard J. Arnott . Morphology and dichotomous vasculature of the leaf of Kingdonia uniflora. Amer. Jour. Bot. 47 (8): 684–698. Illus. 1960.—An intensive study of the nodal anatomy, petiolar vasculature and open dichotomous venation of the leaf of Kingdonia has revealed a type of foliar vascular system of unusual morphological and phylogenetic interest. The vascular supply at the nodal level consists of 4 collateral traces which diverge from a single gap into the sheathing leaf base. This type of nodal anatomy is perhaps primitive, and comparisons are made with the unilacunar nodes and the 2- and 4-parted leaf trace systems characteristic of many angiospermous cotyledons and the foliage leaves of certain woody ranalian genera. The petiole of Kingdonia is vascularized by 2 pairs of bundles which represent the upward continuation of the 4 leaf traces. A transition from an even (4) to an odd (3) number of strands occurs near the point of attachment of the 5, lobed, cuneiform lamina segments to the petiole. Each of the 2 abaxial bundles dichotomizes and the central derivative branches fuse to form a double bundle which enters the base of the median lamina segment. The 2 adaxial petiolar bundles diverge right and left into the bases of the paired lateral segments of the lamina. An analogous type of transition from an even to an odd number of veins occurs in many angiospermous cotyledons which develop a definable mid-vein. But, in Kingdonia, the bundles which enter the bases of the lamina segments give rise to systems of dichotomizing veinlets devoid of “mid-veins.” Although the majority of the terminal veinlets enter the marginal teeth of the lamina segments, “blind” endings, unrelated to the dentations, occur in all the leaves studied. Typically, all of the vein endings in a given lobule of a lamina segment are derived from the same dichotomous vein system. However, in some leaves, a veinlet dichotomizes directly below a sinus and the branches diverge into the marginal regions of 2 separate lobules. The phylogenetic significance of the occurrence of open dichotomous venation in such an herbaceous angiosperm as Kingdonia is briefly discussed. From a purely morphological viewpoint, the Kingdonia type of venation invites direct comparison with the venation of Sphenophyllum, certain ferns or Ginkgo rather than with any of the known reticulate venation patterns of modern angiosperms. Although the foliar venation of Kingdonia may represent the result of evolutionary reversion, the very rare anastomoses which occur seem primitive in type rather than “vestiges” of a former system of closed venation.     

Dpp/BMP transport mechanism is required for wing venation in the sawfly Athalia rosae

The pattern of wing venation varies considerably among different groups of insects and has been used as a means of species-specific identification. However, little is known about how wing venation is established and diversified among insects. The decapentaplegic (Dpp)/bone morphogenetic protein (BMP) signaling pathway plays a critical role in wing vein formation during the pupal stages in Drosophila melanogaster. A key mechanism is BMP transport from the longitudinal veins (LVs) to the posterior crossvein (PCV) by the BMP-binding proteins, short gastrulation (Sog) and twisted gastrulation2/crossveinless (Tsg2/Cv). To investigate whether the BMP transport mechanism is utilized to specify insect wing vein patterns in other than Drosophila, we used the sawfly Athalia rosae as a model, which has distinct venation patterns in the fore- and hindwings. Here, we show that Ar-dpp is ubiquitously expressed in both the fore- and hindwings, but is required for localized BMP signaling that reflects distinct wing vein patterns between the fore- and hindwings. By isolating Ar-tsg/cv in the sawfly, we found that Ar-Tsg/Cv is also required for BMP signaling in wing vein formation and retains the ability to transport Dpp. These data suggest that the BMP transport system is widely used to redistribute Dpp to specify wing venation and may be a basal mechanism underlying diversified wing vein patterns among insects.     

DREWRIA POTOMACENSIS GEN. ET SP. NOV., AN EARLY CRETACEOUS MEMBER OF GNETALES FROM THE POTOMAC GROUP OF VIRGINIA

Drewria potomacensis gen. et sp. nov. from the Lower Cretaceous Potomac Group of Virginia (Zone I, probably Aptian) provides the first definite Mesozoic megafossil record of Gnetales. The stems are slender, display no evidence of secondary growth, and show axillary monopodial branching. Attached leaves are opposite and decussate, borne at swollen nodes, and have clasping sheathing bases. Each leaf is oblong, up to 20 mm long, and has a dense network of longitudinally aligned subepidermal fibers. Leaf venation consists of two, or possibly three pairs of longitudinal parallel veins that form a reticulum at the apex and higher-order crossveins that form apically oriented chevrons. Reproductive structures consist of short, loose spikes that are borne in dichasially arranged groups of three. The dichasia are either terminal or lateral in the axil of a leaf. Reproductive units of the lateral spikes in a dichasium contain seeds, each surrounded by at least one pair of opposite, broadly elliptical or ovate bracts. Characters of D. potomacensis that suggest a gnetalean relationship include the opposite bracts surrounding the seeds, the network of subepidermal foliar fibers, and the distinctive leaf venation, which is very similar to that seen in the cotyledons of extant Welwitschia. Other features consistent with a gnetalean relationship include opposite and decussate leaves, swollen nodes, and the dichasial arrangement of the reproductive spikes. Masses of polyplicate gnetalean pollen comparable to that of extant Welwitschia occur in the same bed as the megafossils. The morphology of D. potomacensis indicates that it was an herb or possibly a shrub. The growth habit of D. potomacensis and associated plants, combined with the sedimentary occurrence of the fossils, indicate that this species and perhaps related taxa were important components of early successional vegetation during the mid-Cretaceous.     

Marginal and laminar hydathode-like structures in the leaves of the desiccation-tolerant angiosperm Myrothamnus flabellifolius Welw.

The fan-shaped leaves of the resurrection plant Myrothamnus flabellifolius Welw. fold during episodes of drought and consequent desiccation of the tissue. The leaf teeth of M. flabellifolius have several features characteristic of hydathodes. Tracheary elements from the three vein endings that converge in each leaf tooth subtend and extend into a cluster of cells significantly smaller than those of the adjacent mesophyll. The stomata overlying this putative epithem are larger than the other stomata on the leaf surface. Crystal violet is absorbed via these stomata in non-transpiring leaves, suggesting that they are water pores. Two to four such water pores occur per hydathode and are readily distinguished in desiccated leaves. Laminar hydathodes apparently also occur in the leaves of M. flabellifolius. Branched vein endings that terminate in short, wide tracheary elements subtend the outer edges of the abaxial leaf ridge, which otherwise lack stomata, and coincide with regions of crystal violet uptake. Guttation could not be induced in M. flabellifolius. However, desiccated leaves readily absorb liquid water through the leaf surface. The use of Calcafluor White to trace the pathway of apoplastic water movement suggests a role for both types of hydathode in foliar water uptake during rehydration while the accumulation of Sulphorhodamine G (indicating solute retrieval from the apoplast) in the epithem of transpiring plants suggests the hydathodes may be a pathway of water loss in the desiccating leaf.     

First report of the family Palaeosetidae (Lepidoptera: Hepialoidea) from mainland China,with description of a new Ogygioses species and its phylogenetic position within the genus

The family Palaeosetidae is newly recorded from mainland China. A new species of the genus Ogygioses, O. maoershana sp. nov., is described from Guangxi Province, China. The adults, male and female genitalia, venation, head and legs structures are illustrated. Some ecological information and a key to all species of the genus are also provided. The phylogenetic position of the new species in the genus Ogygioses is inferred as intermediate between primitive O. luangensis Kristensen and other four derived species based on morphological characters and DNA barcodes sequences.www.zoobank.org/urn:lsid:zoobank.org:pub:D1FD6818-500A-4AE1-9F0B-13CB5A6E1ED4.     

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.     

<Emphasis Type="Italic">Dicroidium bandelii</Emphasis> sp. nov. (corystospermalean foliage) from the Permian of Jordan

Dicroidium bandelii sp. nov. (Corystospermales) is described from the Um Irna Formation (Permian) of the Dead Sea Region, Jordan. The type and only specimen is a compressed frond fragment with cuticle. The new species is unique in showing pinnate frond architecture with elongate, lanceolate pinnae with a dense, near-parallel venation. In situ epifluorescence-microscopic analysis of the cuticle revealed evenly distributed, superficial stomata with differentiated lateral subsidiary cells typical of the genus. Epidermal and cuticular features readily distinguish D. bandelii from some superficially similar species of Ptilozamites or Thinnfeldia. Our report demonstrates that the diversity of Corystospermales in the unusual ‘mixed flora’ of the Permian palaeotropics is still greater than previously thought, and that the Um Irna Formation in particular holds a great potential for future palaeontological research.     

A morpho-anatomical characterisation of Myrothamnus moschatus (Myrothamnaceae) under the aspect of desiccation tolerance

Morpho-anatomical traits of the rarely studied dicotyledonous desiccation-tolerant shrub Myrothamnus moschatus were examined and compared for the first time to Myrothamnus flabellifolius under the aspect of desiccation tolerance. Both species almost exclusively occur on rock outcrops and differ mainly in their geographic range and leaf morphology (fan-shaped in M. flabellifolius, lanceolate in M. moschatus) but have a very similar leaf and wood anatomy, except for the lack of hydathodes in M. moschatus. Both species adopt the parallel leaf venation of monocots, although this is more pronounced in M. moschatus. This provides a mechanical and protective advantage over the net venation pattern of most dicots and facilitates the reversible, drought-induced, accordion-like leaf contraction. The sclerenchyma, as a stabilising tissue, is mainly confined to vascular bundles in leaves of both species. Here, mechanical support seems to be less crucial for survival in long periods of drought than other morpho-anatomical traits (e.g. parallel leaf venation).     

Long Distance Transport in Macrocystis integrifolia: III. MOVEMENT OF THO

Movement of THO and tritium-labeled photoassimilate was studied in intact fronds and frond cuttings of Macrocystis integrifolia following labeling of a mature blade by tritiated water. Both THO and tritium-labeled assimilate moved from the source blade to sink areas at velocities comparable to those recorded earlier for ¹⁴C- and ³²P-labeled compounds. In intact fronds and frond cuttings, THO and tritium-labeled assimilate showed a declining gradient with increasing distance from the source. In the exudate collected from the basal cut end of the frond, there was a marked increase in radioactivity with time in the photoassimilate, but no such gradient was evident for THO. These results support the idea that, although both tritium-labeled assimilate and THO move in the sieve elements, THO is rapidly exchanged with water in the tissues surrounding the sieve elements. Finally, it is shown that THO is transported to the sink and there “unloaded”; indeed, it can move out of the plant itself. The data on velocity and directionality of transport as well as unloading of THO at the sink are discussed, along with computations on specific mass transfer, and favor the idea that Münch's pressure-flow hypothesis is applicable in Macrocystis for long distance translocation of photoassimilates.     

Reconstructing the Pennsylvanian-Age Filicalean Fern Botryopteris tridentata (Felix) Scott

Numerous anatomically preserved fragments of the Middle Pennsylvanian age filicalean fern, Botryopteris tridentata, occur in coal balls collected at the Pittsburgh and Midway Coal Company mine near Baxter Springs, Kansas. Included are the first fertile specimens of the species, evidence of complete vegetative frond architecture, and fronds that are specialized for vegetative propagation. Rhizomes are erect and unbranched, have helical phyllotaxis and short internodes, and typically display an ectophloic solenostele. Fronds are tripinnately compound with lobed pinnules that have open, dichotomous venation. Fertile pinnae or individual pinnules are interspersed among vegetative frond segments and produce sori of annulate sporangia beneath veins on the abaxial pinnule surface. Fertile pinnule lobes are rolled toward the abaxial surface to enclose the sori. Sporangia have a horizontally elongated biseriate annulus located near the short broad stalk and produce tetrahedral-shaped trilete spores with coarse spines. Epiphyllous branches diverge from the stipe or rachis, and some fronds produce only branches. This fern is reconstructed as having short stems. Helically arranged fronds are either pinnately dissected with lobed vegetative pinnules and abaxially rolled fertile pinnules or are specialized for vegetative propagation. The latter functioned as the foliar equivalent of stolons. While some characters of the B. tridentata plant are similar to those of Botryopteris forensis, generitype of the Botryopteridaceae, others are more comparable to those of Psalixochlaena cylindrica, generitype of Psalixochlaenaceae, suggesting the need for reevaluation of systematic relationships among species of the Botryopteridaceae and Psalixochlaenaceae.     

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.     

ONTOGENY OF FOLIAR VENATION IN EUPHORBIA FORBESII

Six species of Euphorbia endemic to the Hawaiian Islands have disjunct veins as a normal component of their foliar anatomy. An ontogenic study of the foliar venation of one of these species, E. forbesii, showed a normal development of the foliar procambium as determined by previous studies of dicotyledonous leaves. The disjunct veinlets are isolated early in the histogenesis of the intersecondary veins when certain procambial cells fail to differentiate into vascular tissue. It appears that these cells develop into normal parenchymatous cells of the ground tissue. It is suggested that these cells are physiologically distinct from the rest of the procambial cells. In no instance was a tracheary element seen which appeared to have arisen independently of the normal procambial reticulum.     

Ediacaran fronds

Leaf-shaped Ediacaran fronds such as Charniodiscus, Charnia, Rangea and Swartpuntia are among the most widespread and readily recognized elements of the soft-bodied Ediacara Biota (late Neoproterozoic). Recent papers have proposed a variety of possible lifestyles for Ediacaran fronds, but functional morphological analysis supports the traditional view that Ediacaran fronds gathered nutrients from the water column through filter feeding or direct nutrient absorption, resulting in the onset of Phanerozoic-style ecological competition complete with denizens occupying distinct tiers in the water column. Within the standard Ediacaran frond morphoshape, four distinct branching architectures have been isolated: Arborea-type branching, characterized by pea-pod like primary branches which house several secondary branches within the protective sheath; Charnia-type branching, resulting in a zigzagging central axis due to the overlapping of alternating sigmoidal primary branches which overly an inferred internal stalk; Rangea-type branching, consisting of petaloids composed of several overlapping primary branches which are self-similar (fractal) over three orders of branching and attached to a straight central stalk; and Swartpuntia-type branching, consisting of petaloids with unornamented tube-like primary branches. Traditional Ediacaran taxonomy has emphasized unity of fronds as a high-level taxon, but the frond morphology most likely represents convergent evolution resulting from competition for nutrients in the water column. It is recommended that any new, higher-order classification schemes highlight the differences in branching architecture as a way of recognizing clades within the Ediacara biota.