Ecological optima and tolerances of Thelypteris limbosperma,Athyrium distentifolium,and Matteuccia struthiopteris along environmental gradients in Western Norway

The distribution and abundance of Thelypteris limbosperma, Athyrium distentifolium, and Matteuccia struthiopteris are modelled statistically in relation to 14 environmental variables along the major climatic, topographic, and edaphic gradients in western Norway. The data are from 624 stands from which measurements or estimates of mean January and mean July temperatures, humidity, altitude, aspect, and slope are available. From 182 of these stands eight soil variables have also been measured. The species responses are quantified by two numerical methods: Gaussian logit regression and weighted averaging (WA) regression. The estimated WA optima suggest that A. distentifolium has an ecological preference for low July and January temperatures, high altitudes, and soils of low-medium pH and base content. The species shows statistically significant Gaussian responses with summer temperature, humidity (= Martonnes humidity index), altitude, slope, aspect, pH, cation exchange capacity, and base saturation with optima of 8.7 °C, 188.9, 1220 m, 28°, 29°, 4.8, 13.77 mEq 100 g dry soil^-1, and 13.4%, respectively. These suggest that the occurrence and relative abundance of A. distentifolium are well predicted by summer temperature, topography, and soil pH and base status. T. limbosperma has WA optima that suggest that it favours moderately high winter and summer temperatures, high humidity, medium altitude, and soils of low pH and base content. It has significant Gaussian responses to summer temperature (optimum =12.6 °C), winter temperature (-1.8 °C), humidity (179.2), altitude (459.5 m), slope (22.5°), and Na (0.7 mg 100 g dry soil^-1). These suggest that climatic factors, altitude, and slope are significant predictors for its occurrence and abundance. M. struthiopteris has high WA optima for summer temperature, pH, Ca, Mg, K, Na, cation exchange capacity (CEC), and base saturation, and a low optima for humidity and winter temperature. Of these, summer temperature (16.0 °C), Ca (63.1 mg 100 g dry soil^-1), Mg (41.0 mg 100 g dry soil^-1), K (23.6 mg 100 g dry soil^-1), Na (5.0 mg 100 g dry soil^-1), CEC (60.7 mEq 100 g dry soil^-1), and base saturation (56.3%) have significant Gaussian logit responses, as do aspect (150.2°) and loss-on-ignition (9.4%). These results suggest that the occurrence and relative abundance of M. struthiopteris are well predicted by high soil base cations, a generally southern aspect, low organic content in the soil, and high July temperatures.     

Quantitative vegetation-environment relationships in west norwegian tall-fern vegetation

The aims of this paper are to detect floristic variation within different types of tall-fern dominated vegetation and to interpret these patterns in terms of environmental variables. Numerical approaches have been applied to a large and varied vegetational data-set with associated environmental data from stands dominated by Athyrium distentifolium, Thelypteris limbosperma , and Matteuccia struthiopteris in different parts of western Norway. The numerical procedures of two-way indicator species analysis, simple discriminant functions, and canonical correspondence analysis have been used, and the strengths and weaknesses of these as tools in discerning vegetational-environmental relationships are discussed. For each of the 96 quadrats investigated, 17 environmental variables were measured. The investigation shows that some of the observed differences in vegetational composition can be explained in terms of relatively simple soil and climatic variables measured for each quadrat. The ferns appear to be ecologically well separated. T. limbosperma-dominaled stands are mainly characterised by low soil fertility, high January temperature, and high humidity. A. distentifolium-dominated stands are associated with low winter temperatures, and M. struthiopteris-dominated stands have high soil fertility and high summer temperatures.     

Seasonal changes in polar lipids in fronds of the ferns Dryopteris filix-mas and Matteuccia struthiopteris

The distribution pattern of total lipids, glyco- and phospholipids, and one betaine lipid (DGTS) in the fronds of the ferns Dryopteris filix-mas and Matteuccia struthiopteris was studied. The lipid composition of the embryo leaflets forming a bud, or treble clef, and that of fully opened leaves changed throughout the growth season. The maximum amount of DGTS in clefs and mature leaves was detected at the beginning of the season. By midsummer, the DGTS content decreased, dropping to zero in the fully opened leaves, and then increased again. The amount of DGTS in the clefs collected in October versus those collected in May was somewhat higher in the case of Dryopteris filix-mas and almost twofold lower in the case of Matteuccia struthiopteris. The ratio between polar lipids contained in the clefs and mature leaves throughout the growth season was determined.     

The effect of enhanced nitrogen on aboveground biomass allocation and nutrient resorption in the fern <Emphasis Type="Italic">Athyrium distentifolium</Emphasis>

We measured aboveground biomass allocation and resorption of nitrogen (N) and phosphorus (P) in the fronds of a winter-deciduous fern Athyrium distentifolium in ambient and N-enhanced treatments. Studies were done in the Moravian-Silesian Beskydy Mts. (Czech Republic) during 2007 and 2008. Athyrium distentifolium formed taller fronds and petioles and allocated more biomass to supporting tissue relative to photosynthetically active leaf tissue in response to N addition. Resorption of P from green fronds was more efficient than N resorption (on average 50% P, 21% N from supporting and 44% P, 24% N from photosynthetic tissues were withdrawn during senescence). The N/P-ratios were higher in photosynthetic tissue (10.8 in 2007 and 13.0 in 2008) in comparison with supporting tissue (5.5 and 7.7, respectively). In N-enhanced treatment, a positive relationship was found between the amount of supporting tissue relative to photosynthetic tissue and resorption of nutrients from photosynthetic tissue. However, higher N availability resulted in a significant decrease of N resorption efficiency in photosynthetic tissue of A. distentifolium.     

Vascular Differentiation in the Shoot Apex of Matteuccia struthiopteris

Initial vascular differentiation is generally considered tooccur in procambium. In ferns, however, a provascular tissueimmediately subjacent to the promeristem has been suggestedas an initial stage within which the procambium is subsequentlyformed. In contrast to this interpretation, a zonation conceptapplied in ferns recognizes a promeristem consisting of severallayers of cells in which no differentiation takes place. Thisstudy demonstrates that the shoot apex of Matteuccia struthiopterishas one cell layer of promeristem. Immediately subjacent tothe promeristem is the provascular tissue surrounding a centralgroup of pith mother cells. The developmental continuity betweenthe provascular tissue and the mature vascular tissue, and betweenthe pith mother cells and the pith, through transitional stages,indicates that the initial differentiation of vascular tissueand pith takes place in this prestelar tissue. The continuityof vascular differentiation in the area confronting young leavesor incipient leaf positions is interrupted by the formationof leaf gap initials. Developing leaves thus begin to exertinfluence on the vascular system at the prestelar stage. Smallprotoxylem elements with helical cell wall thickening, and distinctiveprotophloem elements are present in the leaf traces, but endabruptly near the junction regions of leaf traces to the meristele.Copyright1994, 1999 Academic Press Initial vascular differentiation, provascular tissue, pith mother cells, shoot apex, Matteuccia struthiopteris     

Size and reproduction of Thelypteris limbosperma and Athyrium distentifolium along environmental gradients in Western Norway

The performance of the ferns Thelypteris limbospgrma and Athyrium distentifolium , as expressed by number of son on pinnules and frond size, has been related to environmental gradients. Data have been collected from homogenous fern-dominated stands from different geographical areas in Western Norway. Number of sori produced, which was assumed to indicate fern fertility, was highly variable. There were significant correlations between frond size and fertility, but only 35–46 % of the variation in fertility could be explained by differences in frond size. The highest number of sori was found on plants of medium size, and for A. distentifolium there was a statistically significant reduction in sori number on fronds taller than 125 cm. For the linear relationships between fertility and size, it is suggested that the small X-intercept and slope defines the high competitive ability of A. distentifolium in sites with a long-lasting snow cover. The ferns may dominate in large stands even though their fertility is low or zero, which indicates that they under certain ecological conditions mainly reproduce vegetatively. Both ferns show decreased fertility both toward their altitudinal and lowland distribution limits, but their fertility optima were different. The performances of the two fern species showed highly different responses to geographical and climatical variables. The fertility of T. limbosperma could be predicted by mean July- and January temperatures, humidity and canopy cover, while the fertility of A. distentifolium could not be predicted by any of the site environmental variables investigated.     

Differential Response in the Water Status of Immature and Mature Fronds of the Ostrich Fern (Matteuccia struthiopteris [L.] Todaro) to a Mild Water Stress

Experiments were conducted in growth chambers to examine the effect of a mild water stress (−200 kilopascals polyethylene glycol) on frond elongation and water status of the ostrich fern (Matteuccia struthiopteris [L.] Todaro). Measurements were taken for two days, starting one day after the application of polyethylene glycol. Total water potential in control (well-watered) plants was always significantly higher in immature fronds than in mature fronds. The osmotic potential in mature fronds was always significantly lower (about 800 kilopascals) than in immature fronds in both control and stressed plants. In immature fronds, the stress decreased elongation and total water and pressure potentials, while in mature fronds it increased total water and pressure potentials. The decreases in total and pressure potentials in immature fronds were approximately equal to the increases in mature fronds.     

Reiterative growth in the complex adaptive architecture of the Paleozoic (Pennsylvanian) filicalean fern Kaplanopteris clavata

Reiteration is a widespread component of plant growth whose evolutionary importance in ferns is not recognized widely. We introduce and discuss the growth architecture of Kaplanopteris clavata, a fossil filicalean fern from the Pennsylvanian (ca. 305 million yeas old), focusing on types of reiteration exhibited by this species and on the adaptive and phylogenetic significance of reiteration for ferns in general. Kaplanopteris clavata combines two types of reiterative growth where growth modules are borne on fronds: (1) entire fronds derived from primary pinnae, and (2) epiphyllous plantlets. This combination of reiterative pathways, unique among fossil and living ferns, allowed K. clavata to explore ecospace through an opportunistic combination of scrambling, climbing and epiphytic growth. Kaplanopteris clavata underscores the organographic importance of fronds (as opposed to stems) in the adaptive architecture of ferns, emphasizing functional convergences between the different Baupla̋ne of ferns and angiosperms. This unique combination of reiterative pathways is interpreted as a derived condition illustrating the structural and developmental complexity achieved by some filicaleans during the first major evolutionary radiation of leptosporangiate ferns.     

BIOCHEMISTRY OF FERN SPORE PROTEINS: GLOBULIN STORAGE PROTEINS IN ONOCLEA SENSIBILIS AND OSMUNDA CINNAMOMEA

Onoclea sensibilis was found to contain globulin storage proteins of 2.0S and 11.3S. These globulins, comparable in size and subunit composition to the spore storage proteins characterized in Matteuccia struthiopteris (Templeman, DeMaggio, and Stetler, 1987), declined during imbibition and the initial stages of spore germination. Osmunda cinnamomea, a fern that is only distantly related to Matteuccia, also contained globulin proteins, but these had S values of 5.5 and 11.3. SDS-PAGE analysis revealed extensive differences in banding patterns of the 11.3S protein between Onoclea and Osmunda. This work indicates that while globulin proteins are important storage materials in a variety of ferns they exhibit considerable molecular heterogeneity. The observed heterogeneity in the globulin proteins may be a useful tool to explore evolutionary relationships in the ferns.     

Variations in the population structure and ecology of Matteuccia struthiopteris

The aim of the study was to investigate variation in the structure of a population of Matteuccia struthiopteris (L.) Tod. in N–E Poland depending on its location in a river valley and light availability under tree canopies. Ordination statistical methods and structural equation modeling were used to identify ecological relationships in the population. The analysis showed that optimum growth conditions of M. struthiopteris depend on the canopy cover. As the shading increased, the size of trophophyllous leaves and the production of sporophylls decreased. The relationship between the canopy cover and the optimum growth conditions was best expressed by rootstock diameter and, to a lesser extent, by the number and height of trophophylls. An indirect relationship between the number and height of sporophylls and the canopy cover, by rootstock size and trophophyll quantity, was also identified. This relationship may be caused by the predominance of vegetative forms of reproduction. Based on the correlations between the landscape position vs. rootstock diameter and trophophyll height, a difference in the habitat colonization potential of M. struthiopteris could be detected. Three spatial clusters of M. struthiopteris in the river valley were identified: river bank, river terrace, river valley slopes. These relationships and the distribution of other herbaceous plant species occurring at the plots indicate that the distribution pattern of M. struthiopteris in the study area is determined by diverse environmental conditions, however mainly by light conditions.     

Food preference experiments with captive ptarmigan

Summary

Athyrium distentifolium is found in scattered sites in the Central and Northern Highlands of Scotland. In a few localities in the Central Highlands there is also an endemic variety: A. distentifolium var. flexile. Both types are found together in polymorphic populations with the two distinct forms. A. distentifolium can only grow well with moderate nutrients and in the polymorphic populations A. distentifolium is smaller than elsewhere and frequently infertile. A. distentifolium var. flexile is a smaller plant which is more frequently fertile than A. distentifolium in polymorphic populations and appears particularly well adapted to grow in stressful environments with low nutrient status.     

观赏蕨类引种栽培及其物候期的观察

对中华水韭、福建观音座莲、华南紫萁、西南凤尾蕨、剑叶铁角蕨、翅轴蹄盖蕨、东方荚果蕨、同形鳞毛蕨、圆顶耳蕨等30种蕨类进行了引种栽培和物候期观察.结果表明:(1)引种成活率高;(2)管理粗放;(3)多数种类四季常青,形态优美,具有较高的观赏价值,能够在怀化市安全越冬,值得在亚热带地区开发利用;(4)华南紫萁、小黑桫椤、光蹄盖蕨、长江蹄盖蕨、翅轴蹄盖蕨、三相蕨、同形鳞毛蕨、圆顶耳蕨、镰羽贯众等9种蕨类在展叶时,同时长出孢子囊;(5)多数蕨类的孢子囊在长出后1个月左右发育成熟;(6)在叶开始萌动时引种栽培的蕨类,当年的营养叶萌发期和展叶期都推迟半个月左右,但孢子囊群的出现期和成熟期不受影响;(7)不同蕨类的孢子囊群形成期具有差异性,同种蕨类的孢子囊群形成期具有相对稳定性,因此,可以作为鉴别物种的依据之一。建议在编写、修订《中国植物志》和地方植物志时增加蕨类孢子囊群形成期的描述。     

Leaf development of the ostrich fern Matteuccia struthiopteris (L.) Todaro

The timing of emergence of the three different leaf types of Matteuccia struthiopteris is described from plants sampled over the course of a growing season. Vegetative leaves were first to appear, followed five weeks later by sporophylls and cataphylls. Leaf number and type, and total leaf dry weight per plant were assessed in weekly transects. Vegetative fronds contributed the most to total leaf dry weight, which increased during the first four weeks, and then remained constant for the remainder of the season. Cataphylls, although numerous by the end of the season, contributed little weight. Sporophylls occurred on the widest plants with the most vegetative leaves and greatest leaf weight, whereas cataphylls occurred on most plants except the smallest. Experimentally defoliated plants were re-examined in late summer. Following initial harvest, plants often produced a second smaller set of leaves. These were restricted to vegetative leaves and cataphylls. Ability to reissue leaves, especially vegetative fronds, declined very quickly after the first few weeks in the growing season. Defoliated plants draw on the extensive reservoir of developing leaves which are found on the rhizome, thus possibly diminishing the ability of the plant to withstand regular harvesting of the young fronds for food. Individual leaves were tagged and measured over the growing season. Non-linear regression curves fitted to the growth data for the three types of leaves indicate that growth was described best by a monomolecular growth curve for the vegetative and fertile fronds. Cataphyllar growth could be described equally well by either a monomolecular or a logistic function.     

Characterization of Stelar Initiation in Shoot Apices of Ferns

Procambium is commonly recognized as a vascular meristem inshoot apices of vascular plants. Prestelar tissue comprisingprovascular tissue (PVT) and pith mother cells (PMCs) immediatelysubjacent to the single cell layer of promeristem has been consideredto represent the initial stage of stelar differentiation precedingprocambium and rib meristem in ferns. In addition to characterizationof PVT and PMCs on the basis of cell morphology, cytologicalfeatures and developmental continuity with procambium and ribmeristem, four lines of evidence from studies of shoot apicesof Matteuccia struthiopteris and Osmunda cinnamomea supportthis interpretation of initial differentiation. (1) Differentialstaining by safranin-fast green and crystal violet-erythrosinshows that PVT and PMCs differ in colour reactions from promeristemand resemble procambium and pith meristem, respectively. (2)Comparative ultrastructural study reveals qualitative differencesin the cell membrane system, nuclei, cytoplasm, vacuoles andplastids between promeristem and PVT but similarity of PVT toprocambium. (3) Large droplets of tannins occur in promeristembut not in PVT, PMCs and procambium. (4) Cytochemical studyof the shoot apex of Osmunda shows that carboxylesterase activityis strongly demonstrated in PVT and procambial cells but notin promeristem cells and PMCs. These observations further substantiatethe interpretation that PVT represents initial vascular differentiationand PMCs reflect a commitment to pith development.Copyright1995, 1999 Academic Press Initial vascular differentiation, provascular tissue, differential staining, ultrastructure, tannins, carboxylesterase, shoot apex, Matteuccia struthiopteris, Osmunda cinnamomea     

A tissue culture of the Ostrich fern Matteuccia struthiopteris (L.) Todaro

Multiple bud formation was induced from shoot apices of Matteuccia struthiopteris cultured on semi-solid Knudson's medium supplemented with 10^-5 and 10^-6 M kinetin. The effect of kinetin, naphthaleneacetic acid and gibberellic acid on shoot and root development is discussed and a three-part tissue culture system was devised for micropropagation and rooting.     

A morphometric analysis of Scottish Athyrium distentifolium: a contribution to the BAP

Summary

A morphologically distinct variety of Athyrium distentifolium called A. distentifolium var. flexile has been found only in Scotland. Research was undertaken for aUK Biodiversity Action Plan. To confirm that this taxon has a definitely recognisable morphology, a morphometric analysis was used on the range of characters used to define this variety. It showed that it can be clearly differentiated.     

Geographical variation in frond size,rootstock density,and sexual reproduction in <Emphasis Type="Italic">Matteuccia struthiopteris</Emphasis> populations in Norway

The main aim of this study was to quantify differences in the structure of dense Matteuccia struthiopteris (L) Todaro among geographically separated populations in Norway, and to relate these differences to variation in environmental variables. A stratified random sampling procedure was applied to select 4 m² quadrats restricted to populations dominated (>75% cover) by the fern. This dimorphic fern can produce three types of leaves (fronds): fertile (sporophyll) and sterile (trophophyll), and in addition also intermediate fronds. In all, 178 populations from different parts of the country were analysed, and in total 9,346 rootstocks were measured. Fifteen percent of the rootstocks were fertile, each producing from one to eight sporophylls, and 2.0% had developed intermediate fronds. Rootstocks with five or more sporophylls were mainly found in Southern Norway. Within the plots, there were major variations in density (10–112), number of small (new) ramets (0–58), total number of sporophylls (0–94) and estimated total fern wet-weight (0.33–22.8 kg). All new additions to the populations were connected to a mother plant (ramets), and none were of sexual origin (genets). The main environmental variables associated with the variation in population structure were mean July temperature (highly correlated with altitude and latitude), mean precipitation and density of the canopy cover. The density of small ramets and number of intermediate fronds increased from south to north. The latitudinal variation in sporophyll development and maximum trophophyll height was best explained by a unimodal model. There were statistically significant differences in the population structure between the eight studied geographic areas in Norway.     

Evidence for intergeneric incompatibility in ferns

The archegonial mucilage ofAthyrium filix-femina andA. distentifolium paralyses spermatozoids ofDryopteris filix-mas (and in one caseD. inaequalis) before they penetrate the archegonial venter. The archegonial mucilage ofDryopteris filix-mas has a weak positive chemotactic influence on the spermatozoids of the twoAthyrium species. The spermatozoids ofDryopteris were never observed in the archegonia ofAthyrium. Incompatibility was not observed within and between the twoAthyrium species, withinDryopteris filix-mas or betweenAthyrium filix-femina and twoAsplenium species.Contribution No. 327.     

Effect of Water Stress on Gas Exchange in Fronds of the Ostrich Fern (Matteuccia struthiopteris (L.) Todaro)

Experiments were conducted in a gas exchange system to examinethe effect of a water stress, induced by –200 kPa polyethyleneglycol (PEG), on carbon dioxide and water vapour flux, fronddiffusive resistance, intercellular carbon dioxide concentration,carbon dioxide residual resistance and frond water potentialin the ostrich fern (Matteuccia struthiopteris (L.) Todaro).Measurements were taken 1 d after the application of PEG. Themeasurements were made on young fronds (8 d old) and maturefronds (20–24 d old) at PPFD's (Photosynthetic PhotonFlux Density) from 0–1400 µmol m^–22 s^–1.Water stress decreased the net photosynthesis rate in maturefronds at PPFD's of 210 µmol m^–2 s^–1 or greaterand increased the net photosynthesis rate below 210 µmolm^–2 s^–1 in young fronds. The increase in net photosynthesisin stressed young fronds was associated with a significant reductionin the dark respiration rate. Water stress and decreasing PPFD'sincreased frond diffusive resistance. Carbon dioxide concentrationin the intercellular spaces decreased with increasing frondage and PPFD's up to 200 µmol m^–2 s^–1. Theresidual resistance to carbon dioxide flux was not significantlyaffected by either frond age or water stress. Frond water potentialwas significantly lower in mature fronds than in young fronds. Key words: Matteuccia struthiopteris, Water relations, Photosynthesis, Dark respiration     

Effect of grazing on the population of Matteuccia struthiopteris at the southern limit of its distribution in Europe

We investigated the impact of deer on an isolated marginal population of Matteuccia struthiopteris and on its plant community in the northern Apennines, where in recent decades the species has decreased dramatically. Our experiment was based on a 6‐year before and after control impact (BACI) design, comparing plots with deer grazing and plots where deer were excluded. Exclusion of grazing interacted with years, increasing the size of existing plants and favoring production of sporophylls. The yearly sequence of increase and decline of ramets and sterile leaves followed the same pattern in the two treatments. This indicated that their annual variation in growth depended on the sequence of more and less favorable years. Few and short plants were associated with grazing, whereas tall plants, shrubs, and tree regeneration were associated with fenced plots. The few plants to persist and dominate on heavily grazed areas were Oxalis acetosella and Cardamine species, while the genus Rubus and tree saplings were grazing‐sensitive species in this forest. Tall herbaceous species increased in size in the fenced areas, however, M. struthiopteris drew more advantage in terms of growth of leaves, showing greater cover than other species. Thus, deer grazing is becoming a threat for the survival of M. struthiopteris in southern Europe where it is already threatened in the long term by climate change.