Scaling of Frond Form in Hawaiian Tree Fern Cibotium glaucum: Compliance with Global Trends and Application for Field Estimation

Large‐fronded tree ferns are critical components of many tropical forests. We investigated frond and whole‐plant allometries for Hawaiian keystone species Cibotium glaucum, for prediction and to compare with global scaling relationships. We found that C. glaucum fronds maintain geometric proportionality across a wide range of plant and frond sizes. These relationships result in strong allometries that permit rapid field estimation of frond size from simple linear dimensions. C. glaucum frond allometries complied with intra‐ and interspecific global trends for leaf area versus mass established for much smaller‐leafed species, indicating ‘diminishing returns’ in photosynthetic area per investment in mass for larger fronds. The intraspecific trend was related to declining water content in larger fronds, but not to a significantly larger investment in stipe or rachis relative to lamina. However, C. glaucum complied with the global interspecific trends for greater allocation to support structures in larger leaves. Allometries for frond number and size versus plant height showed that as plants increase in height, frond production and/or retention progressively declines, and the increases of leaf size tend to level off. These frond and whole plant‐level relationships indicate the potential for estimating frond area and mass at landscape scale to enrich studies of forest dynamics.     

Microlepia boluoensis sp. nov. (Dennsteadtiaceae) from Guangdong,China

Microlepia boluoensis Y. Yuan & L. Fu, a new species of the genus Microlepia from Luofu mountain, Boluo county, Guangdong province, China, is described and illustrated. Its gametophytic chromosome number is n =47. Microlepia boluoensis has a similar morphology to M. scyphoformis, M. speluncae and M. hancei, such as creeping rhizomes, bipinnate‐tripinnatified laminas, intramarginal sori, cup‐shaped indusium and trilete spores, but it has a shorter creeping rhizome, larger frond (up to 351 cm), dry chartaceous, longer stipe (up to 139 cm), 23 pairs of primary pinnae, 1–13 sori, and spores with a distinct and raised laesurae.     

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.     

CONTRASTING PATTERNS OF ALLOMETRY AND REALIZED PLASTICITY IN THE SISTER SPECIES MAZZAELLA SPLENDENS AND MAZZAELLA LINEARIS (RHODOPOHYTA)1

Phenotypic differences between the low wave exposure Mazzaella splendens (Setchell et Gardner) Hommersand and the high exposure Mazzaella linearis (Setchell et Gardner) Fredericq could be due to plasticity or genetic differentiation. Common gardens were used to assess their levels of plasticity after describing allometric relationships. As thalli lengthened, stipe development for M. splendens almost ceased even though blades continued to expand, but M. linearis formed a larger stipe before developing a blade that continued to stay narrow at longer thallus lengths. Common gardens demonstrated that M. splendens regrown in the site of M. linearis produced a wider blade than M. splendens regrown in its natural low energy site and that M. linearis regrown in low wave energy either could not form a wider blade or became narrower than thalli from its high energy site. Tetrasporophytes of M. splendens produced a longer and thicker stipe in the high energy site, but the larger M. linearis stipe was not realized because its wider blades made it vulnerable to hydrodynamic removal. Mazzaella splendens therefore had low survivorship in the high wave energy site, and survivors were not long enough to reproduce. Survivorship and reproduction of M. linearis was similar in both environments. Some of the M. splendens and M. linearis characters are plastic, but this plasticity was insufficient for convergence of phenotypes, and blade width plasticity was maladaptive at least for M. splendens. Developmental systems producing the stipe and blade phenotypes of each species have undergone genetic differentiation.     

Alginate content and composition ofMacrocystis pyrifera from New Zealand

Tissue samples ofMacrocystis pyrifera from 2 sites in southern New Zealand and harvested over a period of 12 months were analysed for alginate content and composition (M:G ratio). Plants were divided into three frond classes of different length and each frond was further separated into age categories of blades and stipes (viz young, mature and old blades; mature and old stipes). Within each size class, younger blades had higher alginate content than older blades. Stipes did not show such variation with age. Alginate from younger blades and stipes had higher proportions of mannuronic acid residues than those from old blades and stipes. The range of M:G ratios for age categories of either blades or stipes from longer fronds was greater than those for smaller fronds. Alginate content and M:G ratios of stipes were always higher than for blades. The difference between M:G ratios of blades and stipes was greater for smaller fronds than for longer ones. Differences between collection sites and seasonal trends are also discussed.author for correspondence     

Growth of Ecklonia cava (Laminariales, Phaeophyta) sporophytes transplanted to a locality with different temperature conditions

Transplanting experiments were carried out to determine whether the small type sporophytes with short stipe of Ecklonia cava Kjellman (Laminariales, Phaeophyta) growing in a locality with warm temperatures, change into larger type with a long stipe when transplanted to a locality with cooler temperatures. Juvenile E. cava sporophytes, having a stipe shorter than 5 cm long were collected from Tei in Tosa Bay (southern Japan) (seawater temperature 15–29°C) and transplanted to Nabeta Bay (central Japan) (seawater temperature 13–25°C), where larger type E. cava sporophytes characterized by long stipe (ca 1 m) grow. They were attached to artificial reefs at the sea bottom (9 m depth) in Nabeta Bay to monitor their growth. For comparison, juvenile E. cava sporophytes of almost similar size growing in Nabeta Bay were also transplanted in the same way to the same experimental site. Observations of growth of sporophytes from Tei and Nabeta were carried out monthly for 2 years from November 1995 to October 1997. The transplanted Tei and Nabeta sporophytes showed an increase in stipe length and diameter from winter to spring, whereas almost no increase was observed during summer and autumn. At the end of the study period, the stipe of Nabeta sporophytes reached 25.6 cm in length and 17.0 mm in diameter, whereas that of Tei sporophytes reached 11.1 cm in length and 11.2 mm in diameter. The primary blade length was 16.0 cm in Nabeta sporophytes, whereas it was 5.5 cm in Tei sporophytes. Thus, Tei sporophytes still remained smaller than Nabeta sporophytes even under the same environmental conditions.     

Identification of SCAR marker linking to longer frond length of <Emphasis Type="Italic">Saccharina japonica</Emphasis> (Laminariales,Phaeophyta) using bulked-segregant analysis

To construct a molecular-marker-assisted selection (MAS) system, research was done on identifying molecular markers linking to longer frond length, a crucial selection index in the breeding of the commercially important seaweed Saccharina japonica. An F₂-segregant population of 92 individuals was obtained by crossing two prominent S. japonica strains. Genomic DNA from ten individuals with the longest frond and ten individuals with the shortest frond in the F₂-segregant population were mixed to create two DNA pools for screening polymorphic markers. In bulked-segregant analysis (BSA), out of 100 random amplified polymorphic DNA (RAPD) primers only two produced three polymorphic RAPD markers between the two DNA pools. In conversion of the three RAPD markers into sequence-characterized amplified region (SCAR) markers, only one was successfully converted into a SCAR marker FL-569 linking to the trait of longer frond. Test of the marker FL-569 showed that 80% of the individuals with longest fronds in a wild population and 87.5% of individuals with the longest fronds in an inbred line “Zhongke No. 2” could be detected by FL-569. Additionally, genetic linkage analysis showed that the SCAR marker could be integrated into the reported genetic map and QTL mapping showed that FL-569 linking to qL1-1. The obtained marker FL-569 will be beneficial to MAS in S. japonica breeding.     

Growth and survival rates of large-type sporophytes of Ecklonia cava transplanted to a growth environment with small-type sporophytes

Stipe lengths of sporophytes of Ecklonia cava Kjellman have been reported to be longer along the southeast than southwest coast of the Izu Peninsula, central Japan. Two bays in this region that have natural populations of E. cava, but with different stipe lengths, were chosen for transplant experiments to examine if stipe length was an environmentally controlled trait. Transplant experiments were carried out in order to determine whether large-type sporophytes of E. cava with long stipes growing in Nabeta Bay (southeast Izu Peninsula, Japan) would turn into small-type sporophytes with short stipes when transplanted to Nakagi Bay (southwest Izu Peninsula). Ten juvenile sporophytes of E. cava (stipe length < 5 cm) were collected from Nabeta Bay (large-type habitat) and transplanted to Nakagi Bay (short-type habitat) in December 1995. As a transplant control, ten juvenile sporophytes of E. cava growing in Nakagi Bay were also transplanted to the same artificial reefs. Growth and survival rates of the sporophytes were monitored monthly for 3 y until December 1998. The transplanted sporophytes showed an increase in their stipe length and diameter from winter to spring, whereas almost no increase was observed from summer to autumn. However, the elongation was greater in Nabeta sporophytes than in Nakagi sporophytes. The primary blade length increased mainly from winter to early spring and decreased largely in autumn. Average primary blade lengths were similar in both Nabeta and Nakagi sporophytes from the end of the first year of transplanting. Although ca. 70% of both Nabeta and Nakagi sporophytes survived during the first 2 y after transplantation, no Nakagi sporophytes and only two Nabeta sporophytes survived to the end of the 3 y study period. Despite transplantation to Nakagi Bay, where short sitpes are naturally present, the sporophytes from Nabeta Bay persisted in having longer stipes, which suggests that stipe length is genetically, rather than environmentally, controlled.     

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.     

A new variety ofCortinarius elatior from Japan

Cortinarius elatior var.albipes var. nov., found in the deciduous forest is described and illustrated. It is distinguished from the type variety by having a white stipe and longer spores. The differences betweenCortinarius elatior var.albipes and similar taxa are briefly discussed.     

Effects of nutrient enrichment and herbivory on morphology,reproduction and chemical content of Turbinaria conoides (Phaeophyceae)

The effects of nutrient enrichment and herbivory on resource allocation patterns among morphology, reproduction, and chemical content of the brown alga, Turbinaria conoides (J. Agardh) Kützing were tested in the shallow subtidal zone of the Gulf of Thailand. The field experimental design comprised 36 plots (50 × 50 cm²) with and without herbivores, and two nutrient levels. Cages (uncaged and fully caged plots) were used to exclude herbivorous fishes and two nutrients levels were achieved by experimental enrichment above ambient nutrient concentrations. For morphology of Turbinaria, the maximum length (holdfast to frond apex), the width of the base of the stipe, number of blades, holdfast and stipe diameter were measured. Biomass, reproductive output, and tissue nutrient (carbon: nitrogen: phosphorus) content of T. conoides were examined. Phlorotannin concentrations were examined using a modified Folin‐Ciocalteu method. The results showed that herbivory had no effect on morphology, reproduction, or phlorotannin concentrations. This could be due to the structural and morphological deterrents of alga, which might minimize grazing effects from herbivorous fishes. Nutrient enrichment had no effect on morphology and reproduction of T. conoides, possibly due to low nutrient demand in Turbinaria. However, nutrient enrichment did affect phlorotannin concentrations, as phlorotannins in the enriched plots were lower than the ambient plots. These results support, in part, the carbon–nutrient balance hypothesis that states algae will allocate fewer resources to production of anti‐herbivore chemical compounds when enriched with potential growth‐limiting nutrients.     

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.     

The occurrence of pinhole disease in relation to developmental stage in cultivated <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> (Harvey) Suringar (Phaeophyta) in Korea

Infection rates of pinhole disease in farmed Undaria pinnatifida (Harvey) Suringar were examined between November 2002 and April 2003 at an Undaria farm at Wando on the southwestern coast of Korea. The growth stages of U. pinnatifida can be defined as: a growth phase from December to the end of February, and a decay phase from March to April. There was a significant increase in the level of infection with pinhole disease (measured as the number of pinholes per cm²) between December and April (P < 0.01). Infection progressed from the upper part of the frond in December to the basal part in April. The infection rate of pinhole disease in relation to growth phase on farms over time is limited by the harvest of the seaweed crop at the end of the cultivation period in April. The midrib, sporophyll and stipe showed no signs of infection at any time during the cultivation period. The reduced physiological activity and tissue aging that occurred over time in the sporophytic life phase is the probable cause of the increasing infection rate of the disease.     

Zur Entwicklung und funktionellen Anatomie des Phylloids vonLaminaria hyperborea

The persistent thallus of the phaeophyceanLaminaria hyperborea produces annually a new phylloid, formed by a meristematic zone between stipe and old phylloid. The growing frond is nourished with assimilates from the old one. Young and old phylloid are linked by a collar. Frond and stipe are of very similar anatomical structure. The frond is coated by a one-layer-meristoderm, which forms the external cortex. Using the position of slime ducts as border, the parenchymatous cortex can be subdivided into outer and inner cortex. Between inner cortex and medulla numerous transitions exist. The medulla itself is mainly composed of longitudinally arranged trumpet cells and hyphae. Frond enlargement is caused by the activity of both meristoderm and cortex. Not only cell divisions but also elongation in medulla and inner cortex contribute to growth of the frond. The frond diameter is decreased by this elongation process. Corresponding stretching occurs in the network of slime ducts. Swelling of longitudinal primary walls, accompanied by incorporation of alginate, facilitates the separation of cell strands, and across the medulla hyphae and crosslinks of both inner-cortex-layer cells form. Cells of the inner cortex continously differentiate to elements of the medulla. Trumpet cells within the outer part of the medulla are often branched and connected by hyphae to parenchymatous cells of the inner cortex. Toward the central part of the medulla, trumpet cells elongate and finally attain a length of 1000 µm; they form thick secondary longitudinal walls with ringshaped thickenings, reducing the lumen diameter. Crosswalls and whole cells are often plugged with callose and have, apparently, ceased to translocate.     

Microhabitat,water relations,and photosynthesis of a desert fern,Notholaena parryi

Summary Interrelationships between morphology, microhabitat, water relations, and photosynthesis of a xeric fern, Notholaena parryi D.C. Eat. (Pteridaceae), were examined in the western Colorado desert. In its typical microhabitat rock outcroppings protected N. parryi from direct sunlight and moderated the diurnal variations in air temperature. For example, frond temperature at noon in late winter was 15.3° C, which was 7.3° C cooler than an energy budget simulation predicting frond temperature at an exposed site. The lowest soil water potential leading to daytime stomatal opening was about-1.5 MPa (-15 bars). Rainfall runoff that was channeled to the periphery of the rocks caused ^soil near the fern roots to rise above-1.5 MPa even after light rainfalls, and it remained above-1.5 MPa longer after rainfall than in non-rocky sites.The water potential gradient along the stipe necessary to support the observed rates of transpiration was about-10 MPa m^-1; such a large gradient reflected the small conducting area in the xylem. The water vapor conductance decreased as the frond temperature was raised, an effect that became proportionally greater as the soil dried out. The daytime water-use efficiency (mass CO₂ fixed/mass water transpired) was 0.0058 for a spring day. Individual fronds reached 90% of light saturation for photosynthesis at only 100 Einsteins m^-2 s^-1, a photosynthetically active radiation similar to that from the diffuse sunlight incident on the generally north-facing microhabitat of the fern. Below 50 E m^-2 s^-1 the quantum requirement was 13 Einsteins absorbed/mole CO₂ fixed. The ratio of chlorophyll to P700 was 552, indicating a fairly large photosynthetic unit that is characteristic of plants adapted to shaded habitats. The temperature optimum for net photosynthesis shifted from 13° C in midwinter (mean daily air temperature of 11° C) to 19° C in early fall (air temperature of 23° C).     

Kallymenia crouaniorum (Kallymeniaceae,Rhodophyta), a new red algal species from the Laminaria hyperborea understorey community

In this paper we describe Kallymenia crouaniorum Vergés & Le Gall, sp. nov. (Kallymeniaceae), a new marine red alga from the north-eastern Atlantic Ocean. rbcL and LSU sequences of this species, previously misidentified in the field as Kallymenia reniformis, diverged from those of other Kallymenia species by at least 7.5% and 5.2%, respectively. Kallymenia crouaniorum also has a set of distinctive vegetative and reproductive characteristics, including a deeply lacerate frond, a short stipe, dentate margins, large cortical cells up to 110 μm in diameter, highly refractive stellate medullary cells with arms up to 1000 μm in length, and a monocarpogonial branch system. Molecular phylogenies inferred from rbcL and LSU data indicated, albeit with weak support, that this new species is a sister taxon of a lineage encompassing the generitype K. reniformis, as well as most species of Kallymenia included in the phylogenetic analysis. The main morphological characters that delineate monocarpogonial Kallymenia species are presented. Moreover, after reviewing the literature and several herbarium specimens, we found in the Weber-van Bosse Herbarium a specimen collected at Roscoff in August 1894 with the anatomical characters of the species described here, confirming that this newly described species has in fact been overlooked and is not a recent introduction.     

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.     

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.