Current issue: 54(1)
Under compilation: 54(2)
In an earlier paper of the author it was established that sporal regeneration of bracken fern (Pteridium aquilinum (L.) Kuhn.) is almost entirely connected with fire, and that the size of the bracken fern clones, distinguished by their characteristics, was related to the time of the fire. In previous studies has been found that also sporal regeneration of ground pine (Lycopodium complanatum L., now Diphasiastrum complanatum) is rare. The occurrence of several plant species were studied in relation to bracken fern in the earlier investigation. This paper reports findings concerning ground pine.
When the size-age problem of bracken fern was solved, parallel measurements of ground pine stand on the same site led to the solution of the size-age problem and sporal regeneration of ground pine. The linkage is also valid when the size of ground pine stands was compared to the dates of fires. The ground pine stands are very long-lasting. The stands are fragmented more easily than bracken fern by environmental factors, such as fires, and tend to form large patch clusters with time. Large individual stands reveal the rarity of sporal regeneration of ground pine. The resemblance with bracken fern clones indicate a common factor of regeneration, fire, and a very even spreading rate. Though considerable variation of the colour and structure of ground pine was observed, the circular stands were identical patch by patch.
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In an earlier paper of the author it was established that bracken fern (Pteridium aquilinum (L.) Kuhn.) that the sporal regeneration of the species is almost entirely connected with fire, and that the size of the bracken fern clones, distinguished by their characteristics, was related to the time of the fire. The time of the fire was determined by the samples taken with auger from the trees on the sample plots. A large data on the occurrence of bracken fern was collected around Finland, but only part of it could be used in the previous study, as the trees of the site provided no means to find out it the site had been burned earlier.
In earlier studies, a mass occurrence of sporelings of bracken fern in England has been connected to cities bombed and burned during the World War II. In this study the data of the clones is connected to historical sources and the time of wars in Finland. The comparisons in this study have confirmed the results achieved by comparing the dimensions of the bracken clones to the fire dates. It was found that the wars have caused an increase in the regeneration frequency of bracken fern.
The objective of the present study was to obtain information on the regeneration by spores of bracken fern (Pteridium aquilinum (L.) Kuhn.) as well as the permanence and vegetative spreading of the clones. Prevention of bracken fern, which can hinder forest regeneration, is possible only when the basic facts of the life cycle is known. It would be useful to know if it is probable that bracken fern can become prevalent by spore regeneration, and to what extent it invades forest land by vegetative spreading.
No bracken fern sporelings were found, and the experiments of regeneration by spores were unsuccessful, therefore, the studies were concentrated on the clones. Due to the fire tolerance of the plant, it was expected that the clone material would show gradation according to the periods of fire. The first part of the work concerns individual variation in bracken, the second part concentrates on the data collected from sample plots, which latest fire could be dated from the trees.
The results of the studied clones confirm the correlation between the stands interpreted as clones and the length of the post-fire period. The stands were delimited to clones by their individual characteristics. The spreading on average sites (Vaccinium and Myrtillus type forests) was in average 35.8±2 cm annually. It is concluded that bracken fern regeneration by spores is very rare in Finland and that it has been rarer during the last 50 years compared to the preceding period. This is probably caused by the end of slash-and-burn cultivation and decrease of forest fires in Finland. Bracken fern clones often survive the fires through their rhizomes, and fragments of old clones originating before the fire could be found in the study areas. These clones may be even 300 years old. Especially in its northern areas of distribution, bracken fern is connected with settlement and old slash and burn sites. Regeneration by spores in the northern distribution areas seem to be less than in the southern areas.
Clone stands of bracken fern (Pteridium aquilinum (L.) Kuhn.) wood small reed (Calamagrostis epigeios L.) and lily-of-the-valley (Convallaria majalis L.) are often partly split into two by the road, but often encountered also unilaterally on the roadside in the shape of a semicircle. The unilateral stands can be at times 20–30 m wide and they are sometimes solitary stands of the species. A method to define the age of the solitary stands of six plant species including bracken, wood small reed and lily-of-the-valley was developed in a series of earlier studies.
These stands can be used to define the time the road was built. Clones that are bound by the road unilaterally are younger than the road. If there are several unilateral clones and they are of different sizes, the road is older than the largest clone. When the road is skirted bilaterally only by clones divided by the road, it is younger than the smallest clone. When there are by the road side both unilaterally delimited clones and clones split by the road, the age of the road comes in the range of time determined by the age difference between the largest unilateral and smallest bilateral clone.
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The rate of vegetative spreading in oak fern (Carpogymnia dryopteris) and may-lily (Maianthemum bifolium) was studied by comparing the size of stands formed by individual plants of these species with those of other species growing on the same sites as well as with the time that had elapsed since the last fire on the sites and the age of the tree stand. The average maximum rates of spreading showed to be of similar magnitude in both species, being 12.6 cm/year in oak fern and 12.3 cm/year in may-lily; these values correspond to a radial growth of 6.3 and 6.1 cm year respectively. The maximum radial growth recorded was about 7 cm/year in both species.
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The purpose of this study has been to compile a time-table for the vegetative spreading o the lily-of-the-valley and the wood small-reed. The diameters of the mainly solitary stands of these species have been compared to stand diameters of bracken (Lycopodium clavatum, L. annotium, and L. complanatum), to tree age determined by basal borings, and to times of fires in the site.
Regeneration of the lily-of-the-valley (Convallaria Majalis L.) and the wood small-reed (Calamagrostis epigeios (L.) Roth) from seed is relatively infrequent, and distance between stands may be considerable. Regeneration is more common after fire. The stands of the species are relatively fire resistant, especially on heathlands, where the subsurface parts are safe in the mineral soil. Large stands are generally due to vegetative spreading.
The rate of vegetative spreading of both of the species is practically constant, and about equal, 12.5 cm/year in diameter and 6.2 cm/year in the radial direction. When the actual regeneration time after the fire is not known, the rate of the spreading can be lower than this number.
The present study is a part of larger project into the size and age of certain forest plants. This study seeks to confirm the size-age relationship of Lycopodium clavatum L. and L. annotium L. stands, and the time of sporal regeneration. The stand dimensions were plotted against the size of bracken (Pteridium aquilinum (L.) Kuhn) and ground pine (Lycopodium complanatum L.) stands growing on the same site, and against the age of the timber and the time of fire on the site.
The method based on parallel measurements carried out in burned forests has proved to be suitable for establishing the rate of spreading of stands and their age. The three club-moss species proved to be much alike in their sporal regeneration. They, as well as bracken, regenerate under the condition created by fire. The largest detached patches are often clones. Both Lycopodium clavatum and L. annotium survive fire poorly. Thus, the maximum size of the clones is usually connected with the last forest fire. Pieces of the shoots may survive the fires.
Sporal regeneration of these species also occur without the aid of fire. As also variation in the annual growth of the shoots can be considerable, the stand size of these species is not as good indicator of the date of fire as with Pteridium aquilinum and L. complanatum. The winding appearance of the shoots increases the error in determination of stand age by the stand size further.
The results emphasize the importance of taking into account the time and spread of the plants in the traditional vegetation analysis. A central question is: what is an individual.
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