Fern rhizomes as fodder in Norway

Background

Although ferns are often known under collective names in Norway, e.g. blom, a substantial number of vernacular names for individual fern species are known, in particular for useful or poisonous taxa. In the past, the rhizomes (Norwegian: moldfôr) of selected species were collected for fodder. Only scattered records of such use are available from southern Norway, and the tradition’s core area is found in the two North Norwegian counties of Nordland and Troms, in accordance with the longer winters encountered in the north, frequently leading to fodder shortage in early spring. The tradition extends northeastwards into Finnmark, but is less well documented there. Although numerous sources mention the use of fern rhizomes for fodder, the fern species hiding behind the tradition are incompletely known. This paper aims at reviewing available data in terms of identifyng the species used for fodder, the history and geographical distribution of such use, and other relevant traditions, e.g. the timing and mode of collection, and the way the rhizomes were used.

Methods

The study is based on data extracted from a variety of archival and literature sources; the latter retrived from my database of more than 7500 publications providing information on plant names and plant uses in Norway.

Results

More than 200 individual records mention the use of fern rhizomes for fodder in Norway. Only a fraction of these, typically made by botanist recording data on plant uses, provides information on the identity of the species used. Based on these, Dryopteris filix-mas and Matteuccia struthiopteris stand out as the most important species serving as sources of fern rhizomes for fodder. Locally, Dryopteris expansa was the preferred species, and this taxon may to some extent be overlooked in the records so far available. With a few exceptions, Norwegian folk tradition singles out Athyrium filix-femina as a harmful and poisonous species, causing livestock to go blind and lame, but whether this is true or not, remains unknown; the symptoms are in fact documented elsewhere as a consequence of poisoning due to Dryopteris filix-mas. In coastal north Norway, fern rhizomes were regularly collected for fodder, both in late autumn and early spring, and used to remedy a recurrent shortage of fodder in late winter and spring. Locally, the tradition of collecting fern rhizomes lived on until the 1940’s or 1950’s. Although mainly a tradition of the ethnic Norwegians, it had also been adopted by the farmers belonging to the Finnish and Sámi ethnic minorities.

Conclusion

Fern rhizomes have a long tradition as an additional fodder for livestock in Norway. Preferred species were Matteuccia struthiopteris and Dryopteris filix-mas, locally also Dryopteris expansa. Athyrium filix-femina was considered to be poisonous, and usually avoided.

     

Two new species of Solenostelopteris from the Upper Jurassic Morrison Formation in Wyoming and Utah

Permineralized rhizomes of two new species of Solenostelopteris Kershaw, S. leithii, and S. medlynii, are described from the Upper Jurassic Morrison Formation from the Steiner and Scott's localities north of Greybull, Wyoming, and from Fremont Junction, east of Mt. Ellen, and Mussentuchit Wash sites in Utah. The species are characterized by having a heterogeneous (S. leithii) to homogeneous (S. medlynii) pith, rhizomes with branches (S. leithii), a sclerenchymatous (S. leithii) or parenchymatous (S. medlynii) inner cortex, and both with a parenchymatous middle cortex. The outer cortex of S. leithii is parenchymatous, whereas in S. medlynii, it is composed of round to irregularly shaped, thick-walled cells. The C-shaped leaf traces in S. leithii usually arise single ranked, sometimes two ranked, with short leaf gaps and short internodes. Although the rhizomes of S. medlynii are incomplete and lack leaf traces, they still suggest very long internodes. Because fronds and epidermal trichomes are lacking in both, the affinity of these species to modern taxa, such as members of the Loxsomaceae, remains uncertain.     

Morphology and growth habit of Dicksonites pluckenetii from the Upper Carboniferous of Graissessac (France)

Morphological variability of the vegetative fronds of Dicksonites pluckenetii (SCHLOTHEIM ex STERNBERG) STERZEL is described from new material collected in the Stephanian Basin of Graissessac (Southern Massif Central France). Complete bipartite fronds sometimes still attached to their parent stem allow us to reconstruct the plant as a scrambler/climber. This interpretation is supported by the relatively small slender stems with long internodes and densely covered by short spines, which are also present along the entire length of petioles. A fertile pinna with slightly reduced pinnules bearing up to six radially symmetrical synangia is interpreted as a portion of male frond of the same species; its organization confirms the relationships of Dicksonites with the Callistophytales as was already suggested by features of the female fronds.     

The architecture of a dimorphic clonal fern, Lomagramma guianensis (Aublet) Ching (Dryopteridaceae)

Lomagramma guianensis possesses a dimorphic rhizome and trimorphic fronds. The juvenile genet consists of a slender, sinuous, unbranched, terrestrial rhizome which bears pinnatifid fronds. The adult rhizome form is initiated when the apex becomes scandent. It is stout, exhibiting rhythmic production of pinnate, adult-type fronds, and probably rhythmic growth. The scandent rhizome is ultimately monocarpic, producing fertile acrostichoid fronds and subsequendy dying. Terrestrial runners are produced from proleptic buds at the scandent rhizome base. These are largely similar in morphology to the scandent rhizomes, but display different behaviour. The runners are relatively ephemeral structures which produce no fronds. They branch proleptically from their base to produce scandent rhizomes, and also along their length to give rise to a complex system of runnerS. Senescent runners which have lost contact with their parent scandent rhizome produce reiterative complexes which represent a return to the juvenile form. These reiterated rhizomes behave identically to the juvenile genets, and repeat the entire cycle of development. The architectural history of the clonal pteridophyte Lomagramma is compared to that of angiosperm trees; interesting similarities and differences are found.     

Metals in Tortula muralis from sandstone buildings in an urban agglomeration

Acrocarpous mosses present on stonewalls in urban areas may be used for monitoring purposes. Therefore for this investigation we selected Tortula muralis expecting that this species with an ability to colonise buildings in polluted agglomerations where other species have vanished may provide important information to monitor environmental quality. Concentrations of Cd, Co, Cr, Cu, Fe, Mn, Ni, Pb, V and Zn in T. muralis, in dust deposited on sandstone surfaces, and in the sandstones on which the moss grew were measured. T. muralis appeared to be a good bioindicator of airborne pollution which accumulated especially extremely high Cu, Fe and Zn concentrations reaching values much higher than harmful for plants. There was no relation between metal concentrations in underlying sandstones and T. ruralis. There was a relation between metal concentrations in dust and T. muralis. Cd, Co, Cr, Mn and V concentrations in T. muralis were higher than in dust. The Self-Organizing Feature Map identifying groups of sampling sites with similar concentrations of metals in mosses was able to classify the pollution level by distinguishing groups of highly, medium and less polluted sites. Once trained, SOFM can be applied in ecological investigations and could form a future basis for recognizing e.g. the type of pollution in urban environments by analysing the concentrations of elements in T. muralis.     

Distribution of glycolipids and phospholipids in Pteridium aquilinum

The glycolipids and phospholipids in fronds and rhizomes of Pteridium aquilinum were determined. The total quantity of polar lipid decreased towards the base of the frond, but increased in the storage rhizome. The monogalactosyl diglyceride/digalactosyl diglyceride ratio was 1.8 in the pinnae, 1.0 in the lower petiole and 0.3 in the storage rhizome.     

Seasonal and interspecific variation in the biochemical composition of some British fern species and their effects on Spodoptera littoralis larvae

Fern-feeding insects in Britain are mainly found on mature fronds in late summer. Six fern species (Dtyopteris filix-mas, D. dilatata, D. borreri, Phyllitis scolopendrium, Polyslkhum setiferum and Polypodium vulgare) were analysed for fibre, lignin, cellulose, tannins, cyanogenesis and thiaminase activity in an attempt to determine the biochemical basis for this seasonal pattern of attack. A bioassay was also carried out, using frond material incorporated in the diet of Spodoptera littoralis, to determine the effects of seasonal changes in fern toxins on a non-adapted, insect herbivore. The young fronds of all six species had a high protein content and low fibre-lignocellulose; protein levels decreased and cell wall materials increased as the fronds matured. Tannin concentrations and thiaminase activity showed a less consistent seasonal pattern. Tannins were present in the highest concentration in the young fronds of D. borreri, Polystichum and Polypodium and decreased with season; D. dilatata, Polystichum and Polypodium showed a similar pattern of thiaminase activity. The other species showed no strong seasonality in the levels of either tannins or thiaminase. No cyanogenic activity was detected in any of the fern species. Frond material from all six species decreased survivorship and growth rates of Spodoptera larvae. Larval performance and growth on diets containing fern material from young immature fronds was generally poorer than on diets containing material from mature fronds, collected later in the season. Neither tannin concentrations nor thiaminase activity levels closely corresponded to the observed interspecific and seasonal patterns of larval development and mortality, and other toxins must be involved. It is concluded that ferns are highly toxic to non-adapted herbivorous insects but the effects on adapted species are unknown. The restricted seasonal occurrence of a small number of insect species exhibiting specialized feeding habits suggests, however, that biochemical properties of the ferns determine this pattern of attack and limit the number of species which are able to exploit ferns as food resources.     

Lectins of Dryopteris Adans fern species

Lectin activity of three ferns species (Dryopteris bushiana Fom., D. laeta (Kom.) C. Chr, D. pseudomas (Wollaston) Holub et Pouzar) has been studied. We used hemagglutination reaction albumin and globulin fractions of fronds and rhizomes extracts with rat erythrocytes. The frond extract of D. pseudomas have shown higher activity as compared with other extracts. The lectin activity of D. laeta leaves was absent. The rhizomes of all three fern species could be characterized as high activity. Based on lectin activity the species were placed as follows order: D. buschiana > D. pseudomas > D. laeta. The dependence with lectin activity and elements (Mn, Fe, Cu, Ni, Zn, Pb, Rb, Sr) content were not found.     

安徽石韦属(水龙骨科)一新种

报道了安徽黄山石韦属一新种,即黄山石韦。其与相近种相似石韦的区别主要表现在叶近生,相距1~2mm;能育叶先端不整齐1~2(或更多)回二歧分叉。     

Plants for waste water treatment--effects of heavy metals on the detoxification system of Typha latifolia

Upon treatment with Cd and As cattail (Typha latifolia) showed induced catalase, monodehydroascorbate reductase and ascorbate peroxidase activities in leaves but strong inhibition in rhizomes. Peroxidase activity in leaves of the same plants was inhibited whereas linear increase was detected after Cd treatment in rhizomes.Glutathione S-transferase measurements resulted in identical effects of the trace elements on the substrates CDNB, DCNB, NBC, NBoC, fluorodifen. When GST was assayed with the model substrate DCNB, a different pattern of activity was observed, with strongly increasing activities at increasing HM concentrations. Consequently, to improve the success rates, future phytoremediation plans need to preselect plant species with high antioxidative enzyme activities and an alert GST pattern capable of detoxifying an array of organic xenobiotics.     

Symptoms of Fern Distortion Syndrome Resulting from Inoculation with Opportunistic Endophytic Fluorescent Pseudomonas spp.

Background

Fern Distortion Syndrome (FDS) is a serious disease of Leatherleaf fern (Rumohra adiantiformis). The main symptom of FDS is distortion of fronds, making them unmarketable. Additional symptoms include stunting, irregular sporulation, decreased rhizome diameter, and internal discoloration of rhizomes. We previously reported an association of symptoms with increased endophytic rhizome populations of fluorescent pseudomonads (FPs). The aim of the current study was to determine if FPs from ferns in Costa Rica with typical FDS symptoms would recreate symptoms of FDS.

Methodology and Findings

Greenhouse tests were conducted over a 29-month period. Micro-propagated ferns derived from tissue culture were first grown one year to produce rhizomes. Then, using an 8×9 randomized complete block experimental design, 8 replicate rhizomes were inoculated by dipping into 9 different treatments before planting. Treatments included water without bacteria (control), and four different groups of FPs, each at a two concentrations. The four groups of FPs included one group from healthy ferns without symptoms (another control treatment), two groups isolated from inside rhizomes of symptomatic ferns, and one group isolated from inside roots of symptomatic ferns. Symptoms were assessed 12 and 17 months later, and populations of FPs inside newly formed rhizomes were determined after 17 months. Results showed that inoculation with mixtures of FPs from ferns with FDS symptoms, but not from healthy ferns, recreated the primary symptom of frond deformities and also the secondary symptoms of irregular sporulation, decreased rhizome diameter, and internal discoloration of rhizomes.

Conclusions

These results suggest a model of causation of FDS in which symptoms result from latent infections by multiple species of opportunistic endophytic bacteria containing virulence genes that are expressed when populations inside the plant reach a minimum level.     

EFFECTS OF NITROGEN AND PHOSPHORUS LEVELS,AND FROND-HARVESTING ON ABSORPTION,TRANSLOCATION AND ACCUMULATION OF ARSENIC BY CHINESE BRAKE FERN (PTERIS VITTATA L.)

This hydroponic experiment was conducted to determine the effects of nitrogen (N) and phosphorus (P) levels and frond-harvesting on the effectiveness of arsenic (As)-hyperaccumulator Chinese brake fern (Pteris vittata L.) to remove As from contaminated groundwater collected from south Florida. Three-month old ferns were grown in 38-L plastic tanks (two ferns per tank) containing 30-L of As-contaminated water (130 μg·L^?1 As), which was amended with modified 0.25 strength Hoagland's solution #2. Two N (26 or 52 mg·L^?1) and two P levels (1.2 and 2.4 mg·L^?1) were tested in one experiment, whereas the effect of frond-harvesting was tested in a separate experiment. Initially, N had little effect on plant As removal whereas low P treatment was more effective than high P and As was reduced to <5 μg·L^?1 in 28 d compared to 35 d. For well-established ferns, N and P levels had little effect. Reused fern, with or without harvesting the As-rich fronds, took up arsenic more rapidly so the As concentration in the groundwater declined faster (130 to ～10 μg·L^?1 in 8 h). Regardless of the treatments, most As (85–93%) was located in the aboveground tissue (rhizomes and fronds). Frond As concentrations were higher for non-harvested ferns than for ferns where fronds were partially harvested prior to treatment. Conversely, rhizomes accumulated more arsenic in ferns where fronds had been partially harvested. Low-P treatment coupled with reuse of more established ferns with or without harvesting fronds can be used to effectively remove arsenic from contaminated water using P. vittata     

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.     

New species, newly used names and new ranges of tree ferns (Filicales: Cyatheaceae) in the Neotropics

Two new species are described for the Neotropics, four species are newly accepted, and four species and one variety are reported. The new species are: Cyathea grayumii A. Rojas and C. panamensis A. Rojas. Cyathea alfonsiana L. D. Gómez, C. holdridgeana Nisman & L. D. Gómez, C. onusta H. Christ and C. squarrosa (Rosenst.) Domin are recognized; Cnemidaria coclena Stolze, Cyathea andina (H. Karst.) Domin, C. caracasana var. meridensis (H. Karst.) R. M. Tryon, C. macrosora (Baker) Domin and C. pseudonanna (L. D. Gómez) Lellinger are reported from Costa Rica and Panama.     

Foliar chemistry of balsam fir and red spruce in relation to elevation and the canopy light gradient in the mountains of the northeastern United States

Red spruce (Picea rubens Sarg.) and balsam fir (Abies balsamea[L.] Mill) are the dominant conifer species at treeline in the mountains of the northeastern United States. The objective of this study was to investigate changes in foliar chemistry of these species along both elevational (below, at, and above treeline) and canopy light (sun vs. shade leaves) gradients. Nutrient concentrations (mass basis) did not show any significant (all P>0.05) differences among elevations, although mean concentrations of all macronutrients (N, P, K, Ca, Mg) tended to be higher at low elevation sites compared to high elevation. This result contradicts the traditional view that plants in cold growth environments are adapted to maintain high foliar nutrient concentrations, but it also gives only weak support for the hypothesis that nutrient limitation plays a role in determining treeline location. Foliar concentrations (mass basis) of lignin (both sun and shade needles) and cellulose (sun needles only) decreased sharply and significantly with increasing elevation, but foliar concentrations of hemicellulose did not change with elevation. These results are consistent with the hypothesis that as a result of carbon limitation at high elevation, synthesis of the most expensive fiber constituent (i.e. lignin) is reduced more than that of the least expensive fiber constituent (i.e. hemicellulose). The reduced lignin concentration at high elevation may have implications for nutrient cycling in this ecosystem where cold temperatures limit decomposition rates.     

Dioon califanoi (Zamiaceae), a new species from Mexico

A new Mexican species ofDioon has been found in Oaxaca,Dioon califanoi, characterized by keeled fronds and leaflets inserted obliquely above the rachis. A key to the known species ofDioon is given.     

Open-sea cultivation by transplanting young fronds of the kelp Saccharina latissima

Saccharina latissima is an economically and ecologically important native kelp. As its limited supply from wild stock cannot meet increasing current and future demands, methods for its cultivation in the ocean need to be developed. This kelp is now beginning to be farmed off the Atlantic coast of Spain using a regular method similar to the “forced cultivation” technique used with Asian kelps (kombu). Its cultivation is also a growing enterprise in other European countries. In this study, the open-sea farming of S. latissima using the transplanting method is tested on a commercial scale. This cultivation method has not been studied with kelp species outside Asian waters. The tested method includes the following steps: indoor production of seedlings, pre-culture in greenhouse tanks, and open-sea cultivation by transplanting young fronds. Results demonstrate that open-sea cultivation using transplanted young fronds is a technically and biologically viable method. The mean yield obtained (7.8 kg fresh wt per meter rope equivalent to 45.6 t fresh wt per hectare farm) is satisfactory, considering the low densities of transplanted fronds (25–30 fronds per meter rope). Moreover, these values are comparable to those reported in previous cultivations with this species, as well as in the farming of similar kelps. The transplanting method used in conjunction with the regular cultivation method has valuable practical applications for the commercial farming of S. latissima.     

The significance of rhizome connection of semi-shrub Hedysarum laeve in an Inner Mongolian dune,China

Hedysarum laeve, a rhizomatous clonal semi-shrub, commonly dominates the inland dunes in semiarid areas of northern China. This species propagates vegetatively by extension of horizontal woody rhizomes resulting in programmed reiteration of apical and/or axillary meristems. In this study, the plants were experimentally manipulated by cutting rhizome connections and ¹⁴C-labelling techniques were employed to investigate the ecological significance of rhizome connections within the H. laeve clone. Severance of rhizome connections had a great effect on the performance of young ramets within a clone. Young ramets severed from their parent ramets experienced a significant reduction both in ramet growth and vegetative propagation, as compared with the intact young ramets. Within clonal fragments, consisting of three interconnected ramets including a mother ramet, a daughter ramet and a granddaughter ramet, ¹⁴C-photosynthates from the fed leaves of mother ramets were acropetally transported to all clonal component parts. The ¹⁴C-photosynthate translocation within the clonal fragment provides evidence that the young ramets were supported by their parent ramets. Our results suggest that the woody rhizome connections among the interconnected ramets are ecologically and strategically important for the species to grow in the sand dune habitat.