SPORES OF BOTRYOPTERIS GLOBOSA AND BOTRYOPTERIS AMERICANA FROM THE PENNSYLVANIAN

A study of spores from fertile pinnae of Botryopteris from middle and upper Pennsylvanian coal balls from Iowa, Illinois, and Kansas indicates that there are two distinct species, Botryopteris globosa and B. americana. The organization and attachments of fertile pinnae and the sporangial morphology and dimorphism are identical in the two species. Data are given on fertile pinnae dimensions, attachments of six fertile pinnae, spore counts from individual sporangia, and spore morphology. The ornamentation of B. americana spores is verrucate to rugulate with verrucae fusing to a variable extent to form bars and convolute ridges; B. globosa spores are vermiculate or fossulate to densely rugulate with scattered verrucae. Comparisons are made with B. forensis and a re-interpretation of the spore forms of B. forensis is suggested.     

THE PHYLOGENETIC POSITION OF ANEMIA COLIMENSIS

The unique leaf structure of the genus Anemia with its upright fertile basal pinnae has been interpreted as derived from the typical fern leaf with unmodified basal pinnae. Reported herein for the first time is the leaf morphology of a species, Anemia colimensis Mickel, which shows the most primitive condition known in the genus and clearly confirms the course of evolution. The fertile pinnae are but little modified in size, lamination, and posture. Although A. colimensis has characters of two subgenera, it is shown to be a primitive element in subgen. Anemiorrhiza and not a member of subgen. Coptophyllum as previously reported.     

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.     

INDETERMINATE GROWTH OF LEAVES IN GUAREA (MELIACEAE): A TWIG ANALOGUE

Leaves of seed plants are generally characterized as organs of determinate growth. In this regard, Guarea and related genera seem unusual in that the pinnately compound leaves of these plants contain a bud at their tip from which new pinnae expand from time to time. Previous studies (based upon superficial examinations of leaf-tip buds) have produced contradictory conclusions regarding how long the leaf apex remains meristematic and produces new pinna primordia. In order to determine whether leaf development in Guarea is truly indeterminate, we microscopically examined leaf-tip buds of G. guidonia and G. glabra. In both species, the leaf apex remains meristematic and continues to produce new pinna primordia as the leaf ages. Unexpanded leaves of G. guidonia contained an average of 23 pinna primordia, while the oldest leaves we examined had initiated an average of 44 total pinnae. In G. glabra, unexpanded leaves contained 8 pinnae, whereas an average of 28 pinnae had been initiated on the oldest leaves. These results indicate that leaf development in Guarea is truly indeterminate. Periodic examination of individual intact leaves indicated that the leaves commonly continue their growth for 2 or more years (observed maximum = 51 months). As new leaflets are initiated at the shoot apex (and subsequently expand in rhythmic flushes), older (basal) leaflets may abscise. In addition, the petiole and rachis of the leaf thicken and become woody as a result of the activity of a vascular cambium. Guarea leaves therefore seem to function as the analogue of a typical twig (stem) in general habit as well as in their indeterminate apical growth and secondary thickening.     

AN UNUSUAL BOTRYOPTERID SPORANGIAL AGGREGATION FROM THE MIDDLE PENNSYLVANIAN OF NORTH AMERICA

A new fertile species of Botryopteris (Botryopteridaceae: Filicales) is described from four incomplete Middle Pennsylvanian specimens. Fertile pinnae of B. cratis sp. n. consist of branched frond members bearing numerous globose sporangia. Surrounding the sporangial aggregations are larger sterile frond members (0.5-1.5 mm diam). Fertile pinnae are oval in transverse section and possess an eccentrically developed cortex composed chiefly of fibers. Some frond members show the typical botryopterid xylem configuration with three protoxylem strands. Spherical sporangia are loosely aggregated on the smallest pinnae by short, broad stalks. The annulus is band-like, two cells high, and extends transversely across the lower half of the sporangium for approximately half the circumference. Spores are oval, trilete, verrucate, and covered by a thin separable layer. Sporangium morphology is like that of Botryopteris antiqua, but the spores closely resemble those of B. globosa. The new species is unlike previously described fructifications of Botryopteris in exhibiting a small pinna system which surrounds smaller pinnae bearing sporangia in an aggregation. The new form is considered to be less specialized than previously described globosoid forms because the sporangia are much less crowded. Isolated frond members, believed to belong to the new species, have a large central arm in the pinna xylem trace that resembles the Stephanian taxon B. renaultii. Small stems attached to the adaxial surface of frond members are radial, protostelic, centrarch, and have a three-zoned cortex. The inner cortical zone contains large elongate cells with distinctive layered deposits. Stems are covered with uniseriate multicellular hairs on multicellular bases. Stems compare closely with B. mucilaginosa in histological features.     

Phytoferritin associated with yellowing in leaves ofCocos nucifera (Arecaceae)

Summary Aggregates of phytoferritin particles have been observed by electron microscopy in chloroplasts of leaf pinnae from the coconut palm (Cocos nucifera L.) which have yellowed due to senescence, lethal yellowing disease, scale insect infestation, or physical damage. Phytoferritin was not found in green pinnae from the same trees. We conclude that phytoferritin is a breakdown product associated with disruption of chloroplast structure and photosynthetic activity inCocos nucifera and is not specifically associated with disease.     

Is the large size of the pinna of the ear of the European hare (Lepus europaeus) due to its role in thermoregulation or in anterior capital shock absorption?

Large pinnae are characteristic of the Leporids, and the pinna is known to have a thermoregulatory role. Another role has been hypothesized for the pinna of Lepus spp., as a part of a suspensory system for the greater portion of the head, absorbing shock that might otherwise interfere with vision during high‐speed locomotion. We compared the lengths of the pinnae of adult European hares Lepus europaeus from the source population in the cooler climate of England with those of the introduced population in the warmer climate of Australia, and we compared the lengths of the pinnae of hares that had grown in cooler weather with those that had grown in warmer weather. There were no significant differences between each of the comparisons, indicating that the size of the pinna is not determined by thermoregulatory requirements at rest. We compared the growth in length of the pinnae and the legs with growth in body mass, and growth in the mass of the pinnae with the masses of the head and the eyeballs, and found support for the suspension hypothesis. We suggest that the rapid growth of the pinna is because visual acuity is a function of absolute eye size, not relative eye size, yet juvenile hares are subject to the same predator pressure as adult hares, and equally need to maximize visual acuity while running at high speeds in dim light. We believe that the large size of the pinna is determined by its role in anterior capital suspension, not in thermoregulation. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

PROBABLE ATTACHMENT OF THE DOLEROTHECA CAMPANULUM TO A MYELOXYLON-ALETHOPTERIS TYPE FROND

The discovery of pedunculate specimens of pollen organ Dolerotheca in close association with Myeloxylon-type pinnae and Alethopteris-type pinnules provides evidence for the reconstruction of a petrified medullosan frond and the attachment of the microsporangiate fructification. Specimens of Dolerotheca villosa and D. formosa from Middle and Upper Pennsylvanian age strata of North America are borne on slender peduncles with anatomical features and vascularization identical to those of some level in a Myeloxylon-Alethopteris-type frond. Reconstruction of the frond reveals a large, repeatedly dichotomising organ that bears penultimate pinnae and ultimate pinnules in a pinnate arrangement. Examination of pinnules on the surface of coal ball material indicates that they conform to the compression genus Alethopteris. Identical numbers and arrangement of vascular bundles together with identical anatomical features and multicellular hairs indicate that the Dolerotheca campanulum is borne in the position of a penultimate pinna. A reconstruction of the frond with a proposed polled organ attachment is included.     

CHARLIEA MANZANITANA,N. GEN., N. SP., AND OTHER ENIGMATIC PARALLEL-VEINED FOLIAR FORMS FROM THE UPPER PENNSYLVANIAN OF NEW MEXICO AND TEXAS

Charliea is a new genus (type-species: C. manzanitana), based on pinnately compound leaf material from the richly fossiliferous Virgilian (Upper Pennsylvanian) shales of the Kinney Brick Company quarry near Albuquerque, New Mexico. In several features Charliea resembles Russellites or a zamioid cycad. It has linear-oblong pinnae with broad, oblique attachment and a truncate tip, which is deeply incised to form two to four nearly equal lobes. The venation is simple, parallel, and sparingly dichotomous, each vein ending at the distal margin. The Kinney beds also contain Plagiozamites planchardi, another zamioid form with parallel-veined pinnae, differing from Charliea chiefly in having rounded tips and veins ending in the denticulate margins. An unnamed third form (genus B) in the Kinney beds has long, narrow pinnae with parallel veins and blunt tips; this strongly resembles the Mesozoic conifer Podozamites, but may just as well represent a cycadophyte. Another unnamed taxon (genus A), from an Upper Pennsylvanian deposit in Jack County, Texas, resembles genus B or Russellites in general shape and venation, but the critical distal margins are unknown. In their single-ordered parallel venation, these four foliar types contrast sharply with the two-ordered pinnate venation of most Pennsylvanian fern-like leaves, and seem to foreshadow Mesozoic morphologies. This tendency toward precocious evolution of parallel-veined foliar form in North America is also expressed by a single occurrence of the Asiatic, Permian genus Tingia in the Lower Pennsylvanian of Utah, and by the presence of the predominantly Triassic cycadeoid genus Pterophyllum in the Lower Permian of Texas.     

Effects of MULTIFOLIATE-PINNA, AFILA, TENDRIL-LESS and UNIFOLIATA genes on leafblade architecture in Pisum sativum

In order to dissect the genetic regulation of leafblade morphogenesis, 16 genotypes of pea, constructed by combining the wild-type and mutant alleles of MFP, AF, TL and UNI genes, were quantitatively phenotyped. The morphological features of the three domains of leafblades of four genotypes, unknown earlier, were described. All the genotypes were found to differ in leafblade morphology. It was evident that MFP and TL functions acted as repressor of pinna ramification, in the distal domain. These functions, with and without interaction with UNI, also repressed the ramification of proximal pinnae in the absence of AF function. The expression of MFP and TL required UNI function. AF function was found to control leafblade architecture multifariously. The earlier identified role of AF as a repressor of UNI in the proximal domain was confirmed. Negative control of AF on the UNI-dependent pinna ramification in the distal domain was revealed. It was found that AF establishes a boundary between proximal and distal domains and activates formation of leaflet pinnae in the proximal domain.     

ARCHAEOPTERIS MACILENTA,ANATOMY AND MORPHOLOGY OF ITS FROND

The stem, rachides, and pinnae of Archaeopteris macilenta, formerly considered to be a fern of Devonian age, comprise a branch system in which the ultimate divisions heretofore referred to as pinnules are the leaves. The primary vascular system of the “frond” is a lobed siphonostele from which leaf traces arise in a spiral sequence. The anatomy of the “rachis” and of the “pinnae” is shown to be similar to that of the stem, Callixylon, which bore these “fronds.” Branching, epidermal pattern and stomates are described for the spirally arranged leaves (fertile pinnules). Attachment and dehiscence of sporangia as well as their stomates are reported. Archaeopteris is retained in the Class Progymnospermopsida which includes plants with gymnospermous anatomy and pteridophytic reproduction. It is suggested that Actinopodium, Svalbardia and Siderella are related closely to Archaeopteris and that this group of genera shows evolutionary stages in webbing of leaves and planation of branch systems. The opportunities for ontogenetic studies of the arborescent genus Archaeopteris are pointed out.     

New reconstruction of Weichselia reticulata (Stokes et Webb) Fontaine in Ward emend. Alvin, 1971 based on fertile remains from the middle Albian of Spain

Exceptionally well-preserved specimens from the middle Albian of Spain corresponding to a nearly complete fertile frond and fragments of pinnae containing soral clusters of the tree fern Weichselia reticulata have provided new data about the structure and arrangement of pinnae and their associated fertile reproductive structure. This new material has been compared with the previous studies, and recostructions of this fern and the new data indicate the segregation of vegetative and fertile fronds within the main stipe of this fern.     

Storage ability of overwintering leaves and rhizomes in a semi-evergreen fern, <Emphasis Type="Italic">Dryopteris crassirhizoma</Emphasis> (Dryopteridaceae)

Dryopteris crassirhizoma is a rhizomatous semi-evergreen fern growing in the understory of deciduous forests. Although the top portion of the overwintering leaves began to wither in early winter, intensive senescence occurred in the spring, concurrently with new leaf development. Dry weight comparisons between organs revealed that the rhizome occupied the largest proportion of the total mass, followed by the pinnae. To assess the storage ability of overwintering leaves and the rhizome, seasonal changes in nitrogen content and the dry mass of pinnae and the rhizome were measured. Nitrogen (36.6%) was resorbed from winter-withering pinnae, but not from spring-withering pinnae. In contrast, a similar decrease in dry mass per unit area occurred between winter- and spring-withering pinnae (15%). These results indicate that overwintering leaves serve as a carbohydrate storage organ, but do not serve as a nitrogen storage organ. Nitrogen was not translocated from the rhizome during the early growing season, but translocation did occur in late summer and autumn. The dry mass of the rhizome decreased by 18.4% in spring, at the time of new leaf expansion. The amount of exported dry matter from the rhizome was threefold larger than that from senescent pinnae. Therefore, the rhizome is a major carbohydrate storage organ in this species, although overwintering leaves also act as a carbohydrate storage organ.     

Some ascomycetes (fungi) occurring on tropical ferns

The new combination Eudimeriolum cyathearum is proposed for Dimeriella cyathearum; the fungus is found on pinnae of Cyathea caudata in the Philippine Islands. Dimeriella polypodii is described from scales on pinnae of Polypodium montigenum and P. madrense in Mexico. Bioscypha cyatheae, on pinnae of Cyathea sp. in Costa Rica, is redescribed. Bioscypha pteridicola is described from pinnae of Cnemidaria uleana var. abitaguensis in Colombia. Crocicreas sessilis is described from pinnae of Cyathea divergens var. tuerckheimii in Mexico.     

A quantitative comparison of foliage development among allopatric ferns,Dryopteris crassirhizoma,D. coreano-montana andD. filix-mas

A quantitative comparison was conducted on the foliage development during sporophyte development of three allopatric ferns in cool temperate and subalpine regions of Hokkaido and Tirol, European Alps. The foliage development ofDryopteris crassirhizoma, D. coreano-montana andD. filix-mas was quantitatively described by the leaf development (NV, number of veins); NV correlates the leaf-shape complexity from a circle (DI, L/2(3.14×S)^1/2). Nearly similar patterns were detected on frequency distribution of fertile leaves, fertility increase and number of leaves in threeDryopteris ferns which exhibit funnel-shaped foliage arrangements in mature sporophyte. No difference was found in number of leaves, maximum NV, fertility rate and leaf-shape parameters among three ferns. A positive difference was found only on changes in order of pinnae with maximum number of costa branches (NVP) and the DI of outline of pinnae betweenD. crassirhizoma andD. filix-mas. These allopatricDryopteris ferns seem to have a similar foliage structure, in spite of some sympatricDryopteris ferns capable of producing putative hybrids (D. austriaca andD. amurensis; D. tokyoensis andD. monticola) having different foliage structures in Hokkaido. Contribution No. 3346 from the Institute of Low Temperature Science, Hokkaido University.     

Zur Bewertung der Blattdornen vonHalimodendron halodendron (Fabaceae)

Halimodendron halodendron is a spiny shrub of north-western Asia. The spines are formed by the rachises of the paripinnate leaves. After abscission of pinnae the rigid rachises remain for several years. In culture these rachis spines do not develop fully.

Herrn Univ.-Prof. Dr.Walter Leinfellner zum 70. Geburtstag gewidmet.     

STENOKOLEOS BIFIDUS SP. N. IN THE UPPER DEVONIAN OF NEW YORK STATE

Stenokoleos is a genus for petrified axes from the Mississippian New Albany Shale to which an Upper Devonian occurrence in New York is added. Two orders of branching were known and the plant was thought to be related to coenopterid ferns. The new petrified axes from New York reveal three orders of branching. A pair of rachides emerges from one side of the stem at each node. Their position alternates at successive nodes (distichous). Each rachis bears alternately arranged pinnae. The shape of the xylem strand and the number of protoxylem areas are variable. Traces to the pairs of rachides arise either as two separate strands or as a single strand that is presumed to divide while still within the cortex of the stem. Traces to pinnae are ellipsoid or clepsydroid. Tracheids are scalariform and uni- or biseriate, circular-bordered pitted. Peripheral loops are present in all orders of branches. Protoxylem strands are numerous and maturation is mesarch. Cortex is parenchymatous where it is preserved but outer cortex is missing. Stenokoleos and Reimanniopsis are placed in a new family, Stenokoleaceae. This is classified as Incertae Sedis among Pterophytina in Tracheophyta. It is suggested that the plant is related more closely to the Mississippian pteridosperms Tristichia and Tetrastichia than to the coenopterid ferns.     

Surface structure and anatomical aspects of Chinese brake fern (Pteris vittata; Pteridaceae)

The objective of this study was to describe a wide spectrum of surface structural and anatomical details of the Chinese brake fern (Pteris vittata) using scanning electron microscopy (SEM). SEM revealed that the epidermal cells of the pinnae were elongated with raised periclinal and sinuous anticlinal walls. The pinnae were hypostomatous with randomly scattered anomocytic stomatal complexes positioned at the same level as the epidermis. Stomates were large and elliptical (27.4 μm × 10.2 μm). Cross sections from the central regions of the rachis and the stipe revealed V- and U-shaped vascular bundles, respectively. In each vascular bundle, the xylem strands were sea-horse shaped (hippocampus). In contrast, the pinnae possessed a triangular vascular bundle with uniform mesophyll organization comprising of homogenous lobed parenchyma cells. The indumentum consisted of trichomes and scales, which formed various types of vestiture. Trichomes were borne only on the pinnae and scales on the rachis and stipe. The roots developed a dense network of long root hairs averaging 244 μm long, and the xylem consisted of tracheids with scalariform pitting. Sori were submarginal; continuous along both margins of the pinna and were covered with a false indusium. The sporangia were oblong with a short thick stalk and the annulus was positioned vertically resulting in transverse dehiscence of the sporangium. The paraphyses were uniseriate, unbranched, septate and found to be intermixed with the sporangia. The exine of the globose spores was adorned with thick reticulum in which the areoles contained round tubercles. This study describes surface features in detail, which is essential to studies examining the issue of whether morphological characteristics are related to arsenic hyperaccumulation inP. vittata.     

The external ear morphology and presence of tragi in Australian marsupials

Multiple studies have described the anatomy and function of the external ear (pinna) of bats, and other placental mammals, however, studies of marsupial pinna are largely absent. In bats, the tragus appears to be especially important for locating and capturing insect prey. In this study, we aimed to investigate the pinnae of Australian marsupials, with a focus on the presence/absence of tragi and how they may relate to diet. We investigated 23 Australian marsupial species with varying diets. The pinnae measurements (scapha width, scapha length) and tragi (where present) were measured. The interaural distance and body length were also recorded for each individual. Results indicated that all nectarivorous, carnivorous, and insectivorous species had tragi with the exception of the insectivorous striped possum (Dactylopsila trivirgata), numbat (Myrmecobius fasciatus), and nectarivorous sugar glider (Petaurus breviceps). No herbivorous or omnivorous species had tragi. Based on the findings in this study, and those conducted on placental mammals, we suggest marsupials use tragi in a similar way to placentals to locate and target insectivorous prey. The Tasmanian devil (Sarcophilus harrisii) displayed the largest interaural distance that likely aids in better localization and origin of noise associated with prey detection. In contrast, the smallest interaural distance was exhibited by a macropod. Previous studies have suggested the hearing of macropods is especially adapted to detect warnings of predators made by conspecifics. While the data in this study demonstrate a diversity in pinnae among marsupials, including presence and absence of tragi, it suggests that there is a correlation between pinna structure and diet choice among marsupials. A future study should investigate a larger number of individuals and species and include marsupials from Papua New Guinea, and Central and South America as a comparison.     

Quantitative comparison of foliage development amongDryopteris monticola,D. tokyoensis and a putative hybrid,D. kominatoensis in Northern Japan

Developmental leaf architecture was quantitatively described in terms of measurements of various parameters on leaf blade from different size of sporophytes inDryopteris monticola, D. tokyoensis and a putative hybrid,D. kominatoensis in the natural site of Hokkaido, to compare the ontogenetic differentiation in foliage structure among allied ferns. The morphological stage of leaf and sporophyte was tentatively quantified by the number of midrib branches of the leaf (NV, number of veins), which exhibited a significant correlation to the leaf-shape complexity from a circle (DI=marginal length/2×(3.14×square)^1/2) of leaf blade. D. kominatoensis showed intermediate values between others in following characters; DI increase, maximum NV (also blade length), maximum number of costa branches of pinnae (NVMP), number of costa branches of the lowest pinna (NVLP), difference between NVMP and NVLP (NVMP-NVLP), during heteroblastic leaf development. A larger number of leaves per sporophyte was found inD. kominatoensis than in others. The fertility rate (%) and initiation of fertility (IF) in the relative developmental stage (RDS) ofD. kominatoensis shifted to that ofD. tokyoensis, while the order of pinnae with NVMP shifted to that ofD. monticola. Even in the intermediate characters inD. kominatoensis, slight shifts in characters to those of putative parents were found during heteroblastic leaf development. Contribution No. 3145 from the Institute of Low Temperature Science, Hokkaido University.