Phytometric assessment of alder seedling establishment in fen and bog: implications for forest expansion mechanisms in mire ecosystems

Aims

Rapid forest expansion as a result of anthropogenic activities has been observed in many mires. However, it is unclear which environmental factors are driving this expansion because there have been no systematic investigations into mire-specific tree seedling establishment. This study investigated factors affecting the establishment of common alder (Alnus japonica) in a mire.

Methods

We performed seed sowing and seedling transplantation field experiments to examine the factors influencing germination rate, seedling survival, and seedling growth of A. japonica.

Results

Germination rate and seedling survival period decreased with increasing water level, and seedling dry weight was reduced at pH <6.0. Germination rate was also lower in the fen, whereas seedling dry weight was lower in the bog, which could be reasonably explained by the higher water level in the fen and the lower pH in the bog.

Conclusions

Our results showed that germination and seedling growth in the fen and bog were each inhibited by different mire-specific conditions: high water level and low pH, respectively. Therefore, seedling establishment could be improved by either lowering the water level in fens, to increase germination rate and survival, or raising the pH in bogs, to increase growth.     

Climate change and land use threats to species of aapa mires,an EU priority habitat

Aapa mires are EU priority habitats that harbour unique biodiversity values but face increasing global change threats. Here, we investigate the exposure of red-listed aapa mire species inhabiting fen and flark fen habitats to the impacts of land use and climate change. Climate change-based threats were assessed across the aapa mire zone of Finland based on climate velocities (a metric describing the speed and direction of climate movement) measured for mean January temperature (T_Jan), growing degree days (GDD5) and mean annual water balance (WAB). Land use threats were assessed based on the cover of drainage ditches and three other adverse land use types around the species occurrences. Our results suggest that rapid changes in T_Jan may alter winter thermal conditions and thereby also species performance, particularly in the northernmost part of the aapa mire zone, where the most valuable concentrations of red-listed species are situated. The land use and GDD5 threats are highest in the southern regions where the red-listed aapa mire species occurrences are sparser but face severe risks to their persistence. In the central part of the aapa mire zone, a number of valuable aapa mires with red-listed species are exposed to both intermediately high T_Jan and GDD5 velocities and a spatially varying amount of ditching. Three conservation approaches to support the persistence of red-listed aapa mire species: (i) restoration, (ii) establishment of new protected areas, and (iii) monitoring of the key habitats, should be flexibly and complementarily applied to the preservation of aapa mires subject to accelerating climate change.     

Crown development in tropical rain forest trees in gaps and understorey

Emergence and establishment of Tsuga sieboldii seedlings were monitored for six years in an old-growth mixed forest in Japan, to clarify the reason why adult T. sieboldii trees are distributed only on convex slopes. T. sieboldii seedlings germinated on exposed soils without broad-leaved litter cover on both convex and concave slopes, but they died out from concave slope during the first two years after germination. Along a gradient from convex to concave slope, litter cover became higher and surface soil erosion became severer, and T. sieboldii seedlings died at the plots with severe surface soil erosion. Other environmental variables, such as relative illuminance, vegetation cover of seedling layer and soil water potential were not significantly different between convex and concave slopes. We concluded that the restricted seedling establishment of T. sieboldii to convex slope is determined by stability of surface soils and broad-leaved litter cover.     

Litter controls plant community composition in a northern fen

The accumulation of litter or thatch can affect plant community composition by affecting the temperature, nutrient availability, and light availability of the soil environment, thereby forming a potentially important linkage between recent productivity and current ecosystem processes. To investigate the importance of litter on a fen peatland plant community, we conducted a litter addition and removal experiment in a fen in northern Minnesota, USA, between 1998 and 2001. The addition of litter had little effect on fen plant community composition or microenvironmental variables, despite a two-fold increase in litter mass compared to control plots. However, the removal of litter dramatically increased cover of Rhynchospora fusca and R. alba , and reduced cover of Carex exilis . Litter removal also increased availability of light and soil temperature and increased the phosphorus content of aboveground plant biomass. Our results indicate that litter is an important control of plant community composition in this northern fen.     

Multivariate path analysis of the relationships between seedling regeneration and environmental factors beneath a dwarf bamboo understory

Seedling emergence and establishment are fragile processes that determine the direction and structure of forest succession and regeneration. However, seedling emergence and establishment are easily affected by biotic and abiotic (environmental) factors. A dense and expanding understory of dwarf bamboo is one such important factor that can seriously hinder the seedling regeneration. We conducted a field experiment to investigate the emergence and establishment of canopy tree seedlings under artificially controlled densities of dwarf bamboo. We found that understory dwarf bamboo obstructed seedling emergence but reduced the death of seedlings. Although understory dwarf bamboo reduced the median retention time of seedlings, dense bamboo increased the mean survival time of seedlings. Our results suggest that understory dwarf bamboo has multiple selectivities for tree seedling emergence and establishment: high‐density dwarf bamboo was beneficial to evergreen species but lower‐density of bamboo was conducive to the survival of deciduous species, it means the dwarf bamboo potentially alters successional trajectories of forest communities. Path analysis revealed that the most important factors affecting tree seedling emergence and death were the abundance of seeds in the seed bank and the density of emerged seedlings, and that the soil temperature promoted seedling emergence but increased seedling death, the thickness of litter limited seedling emergence, and the leaf area index of the bamboo canopy limited seedling death. The present study suggests that dwarf bamboo can directly alter the microenvironment, significantly reducing light levels and soil temperature but increasing the thickness of litter and soil humus, thereby indirectly impacting the regeneration of tree seedlings. Our results indicate that various factors affected seedling emergence, and there were complex indirect relationships among these factors. In general, biological factors had a stronger influence on tree seedling regeneration than environmental factors.     

Limiting mechanisms in the regeneration of the Chilean matorral – Experiments on seedling establishment in burned and cleared mesic sites

The Chilean matorral is characterized by multispecific shrub clumps in dry areas but has a continuous canopy in wetter sites. It has been hypothesized that this difference is due to easier recolonization of open patches by shrub seedlings under more mesic conditions. Within the mesic range of the matorral we designed a field experiment to compare shrub seedling emergence, growth, and survival under the closed canopy of a secondary forest versus three types of open patches: burned, cleared of shrubs but with a herbaceous layer present, and clear without a herbaceous layer. After the first summer, survival of Quillaja saponaria seedlings was 75% in the burned site, 30% in the cleared patch without herbaceous vegetation, and 15% with herbaceous vegetation present, whereas there was 0% survival under the secondary forest canopy. After eight years, the percentages had dropped to: 22%, 12% and 3%, respectively. These results contrast strongly with the seedling establishment patterns in drier areas of the matorral where early seedling survival is higher under the shade of large shrubs. In the experimental mesic sites, seedlings did best on the burned site, not only in terms of survival, but also in terms of growth. After one year, seedling mean height was 10.2 cm in the burned site, whereas 3.8 cm and 5.3 cm in the cleared patches without and with herbaceous respectively. After eight years, mean height differences between treatments had increased further: 147.7 cm in the burned site, 40.3 cm in the cleared patch without herbaceous cover and 13 cm in the cleared patch with herbs. Our results indicate that the facilitative effect of nurse shrubs on seedling establishment found in dry ranges of the matorral is less important in more mesic sites. This difference may explain the continuous shrub cover in relatively mesic areas as opposed to the characteristic patchy structure of the matorral in its drier range.     

Seed deposition in drift lines: Opportunity or hazard for species establishment?

Arrival at suitable sites for germination and establishment is crucial for hydrochorous plants. Seed deposition in riparian landscapes occurs mainly within litter in drift lines. We investigated whether conditions within drift lines are suitable for seed survival and germination and whether drift lines act as seed sources or sinks for species diversity in riparian fen grasslands.We analysed drift litter in a small river valley in Northern Germany (Upper Eider valley) after typical late winter floods in 2000, 2002 and 2004. The content of viable seeds in drift lines was studied via a germination trial with drift litter samples taken from five fen grassland sites representative of the vegetation of floodplain. Further, seedling recruitment of both abandoned and moderately grazed fen grasslands affected by drift line deposition was compared with equivalent sites without drift lines. Additionally, the decline of viable seeds after the deposition of drift litter was studied. Finally, the relative importance of seed-containing drift litter and microsite availability for seedling recruitment was investigated in a field experiment with the factors disturbance (mowing, large gaps, small gaps, undisturbed control) and addition of (seed-containing) drift litter (yes, no) in an abandoned riparian fen.Seedlings which recruited from drift litter collected at different sites differed in species composition. A nearly 50% reduction of the number of viable seeds in drift litter was observed already during 2 weeks after deposition. Naturally deposited drift lines had a positive effect on the number of seedlings, although an increase of seedling recruitment in areas with drift litter was only observed at the grazed site. Adding drift litter reduced the number of seedlings in the field experiment, while creating gaps enhanced seedling recruitment.These results indicate that in riparian fen grasslands, drift lines can affect species diversity either negatively by hampering germination and/or establishment or positively by increasing seed availability. As the studied river system is comparatively species poor and flooding extension is limited, the role of drift lines as seed source might be of minor importance in the study area.     

Hydrological landscape settings of base‐rich fen mires and fen meadows: an overview

Question: Why do similar fen meadow communities occur in different landscapes? How does the hydrological system sustain base‐rich fen mires and fen meadows? Location: Interdunal wetlands and heathland pools in The Netherlands, percolation mires in Germany, Poland, and Siberia, and calcareous spring fens in the High Tatra, Slovakia. Methods: This review presents an overview of the hydrological conditions of fen mires and fen meadows that are highly valued in nature conservation due to their high biodiversity and the occurrence of many Red List species. Fen types covered in this review include: (1) small hydrological systems in young calcareous dune areas, and (2) small hydrological systems in decalcified old cover sand areas in The Netherlands; (3) large hydrological systems in river valleys in Central‐Europe and western‐Siberia, and (4) large hydrological systems of small calcareous spring fens with active precipitation of travertine in mountain areas of Slovakia. Results: Different landscape types can sustain similar nutrient poor and base‐rich habitats required by endangered fen meadow species. The hydrological systems of these landscapes are very different in size, but their ground water flow pattern is remarkably similar. Paleoecological research showed that travertine forming fen vegetation types persisted in German lowland percolation mires from 6000 to 3000 BP. Similar vegetation types can still be found in small mountain mires in the Slovak Republic. Small pools in such mires form a cascade of surface water bodies that stimulate travertine formation in various ways. Travertine deposition prevents acidification of the mire and sustains populations of basiphilous species that elsewhere in Europe are highly endangered. Conclusion: Very different hydrological landscape settings can maintain a regular flow of groundwater through the top soil generating similar base‐rich site conditions. This is why some fen species occur in very different landscape types, ranging from mineral interdunal wetlands to mountain mires.     

Abiotic constraints on the establishment of Quercus seedlings in grassland

High evaporative demand and periodic drought characterize the growing season in midwestern grasslands relative to deciduous forests of the eastern US, and predicted climatic changes suggest that these climatic extremes may be exacerbated. Despite this less than optimal environment for tree seedling establishment, deciduous trees have expanded into adjacent tallgrass prairie within the last century leading to a dramatic land cover change. In order to determine the role of light and temperature on seedling establishment, we assessed carbon and water relations and aboveground growth of first‐year Quercus macrocarpa seedlings exposed to one of three conditions: (1) intact tallgrass prairie communities (control), (2) aboveground herbaceous biomass removed (grass removal), and (3) shade plus biomass removal to reduce light (PFD) to levels typical of the grassland‐forest ecotone (shade). In the 2000 growing season, precipitation was 35% below the long‐term average, which had a significant negative effect on oak seedling carbon gain at midseason (photosynthesis declined to 10% of maximum rates). However, net photosynthesis and stomatal conductance in the shade treatment was ca. 2.5 and 1.5 times greater, respectively, than in control treatment seedlings during this drought. During this period, leaf and air temperatures in control seedlings were similar whereas leaf temperatures in the shade treatment remained below air temperature. A late‐season recovery period, coincident with decreased air temperatures, resulted in increased net photosynthesis for all seedlings. Increased photosynthetic rates and water relations in shaded seedlings compared to seedlings in full sun suggest that, at least in dry years, high light and temperature may negatively impact oak seedling performance. However, high survival rates for all seedlings indicate that Q. macrocarpa seedlings are capable of tolerating both present‐day and future climatic extremes. Unless historic fire regimes are restored, forest expansion and land cover change are likely to continue.     

Optimal habitat conditions for the globally threatened Aquatic Warbler Acrocephalus paludicola in eastern Poland and their implications for fen management

To identify optimal habitat for the Aquatic Warbler Acrocephalus paludicola, a flagship species of fen mires, we related numbers of singing males to habitat variables in all core breeding sites in eastern Poland. The density of male Aquatic Warblers increased with increasing ground cover by water and mosses and litter layer height, and was highest where vegetation was 60–90 cm tall. Male densities also increased with the biomass of arthropods > 10 mm length, estimated by sweep netting, and with the abundance of spiders, estimated by pan trapping. We suggest that habitat management should take into account species‐specific morphological adaptations, nest safety and arthropod productivity. Prevention of vegetation succession is a conservation priority for open fen mires. However, modern management practices to achieve this, especially mowing using tracked vehicles, should be evaluated and optimized to ensure that such practices do not adversely affect the long‐term development of moss cover and litter structure.     

浙江天童常绿阔叶林地被层的研究

 对天童常绿阔叶林地被层植物种类组成及群落学作用进行了初步研究。研究表明：组成地被层计有维管植物44科78属114种。蕨类植物优势度最大,里白占绝对优势。39种草木植物,以世界分布科和地面芽植物为主,常绿种类占84.6％,未见亚洲和北美洲间断分布的地理成分。乔灌木幼苗51种,80%的苗量集中于4～6个种中,优势种突出。藤本植物24种,草质藤本占25％,落叶种类42％。地面苔藓植物(22种)与枯枝落叶构成镶嵌分布,发育较成熟的群落中苔藓种类多,盖度大。里白及其枯死物形成层层交错重叠的地被层,阻碍凋落物进入地表,减弱光照,使近地表光照长期低于幼苗光补偿点,抑制了乔灌木优势种的更新。凋落物阻碍种子下移至半分解层,栲树种子下移率为17％至22.3％,冬季种子的动物损耗率平均为98.2％,凋落物中种子的平均腐烂率为28.2％,表层保存下来的栲树种子因缺水萌发速度慢且成活率低,下层萌发速度虽快,但腐烂率高,栲树天然更新幼苗主要来源于草窗。     

Impact of rewetting on the vegetation of a cut‐away peatland

Abstract. We tested whether rewetting improved environmental conditions during peatland restoration and promoted colonization and development of mire vegetation. Vegetation change was monitored in a cut‐away peatland one year before, and four years after, rewetting. Colonizers before rewetting were perennials, mostly typical of hummocks or bare peat surfaces. The main variation in vegetation was related to variation in the amounts of major nutrients and water table level. The wettest site with the highest nutrient level had the highest total vegetation cover and diversity, as well as some species typical of wet minerotrophic mires. Raising the water table level above, or close to, the soil surface promoted development of wet minerotrophic vegetation. Diversity initially decreased because of the disappearance of hummock vegetation but started to recover in the third year. Eriophorum vaginatum and Carex rostrata were both favoured, and bryophytes typical of wet habitats colonized the site. Moderate rewetting promoted the development of Eriophorum vaginatum seedlings and an increase in the cover of tussocks. Bryophytes typical of disturbed peat surfaces spread. In the control site development continued slowly towards closed hummock vegetation. The study showed that raising the water level to, or above, soil surface promotes conditions wet enough for a rapid succession towards closed mire vegetation.     

Young restored forests increase seedling recruitment in abandoned pastures in the Southern Atlantic rainforest

Planting seedlings is a common technique for abandoned pastures restoration in the tropics, supposedly by increasing the seedling recruitment and accelerating succession. In this study we evaluated the role of a young restored forest (one year old) in enhancing seedling establishment from two sources (seed rain and seed bank), in the Atlantic Rainforest region in Southern Brazil. We compared abandoned pasture, young restored forest and old-growth forest with respect to the seedlings recruited from different sources, by monitoring 40 permanent plots (0.5 m x 0.5 m) over 20 months. From the three studied areas a total of 392 seedlings of 53 species were recruited. Species were mainly herbaceous (85%), pioneers (88%), zoochorous (51%) and small-seeded species (60%). Seedling recruitment from the seed bank (density and species richness) was higher and dominated by herbaceous species in the abandoned pasture and in the young restored forest; on the other hand, the recruitment of woody species from seed rain was more pronounced in the old-growth forest. The young restored forest increased the species richness of woody seedlings recruitment from the seed bank (two-fold) and from seed rain (three-fold) compared to the abandoned pasture. Also, the seedling density in young restored forest was still higher than abandoned pastures (seed bank: four times; seed rain: ten times). Our results show that even young restored areas enhance the establishment of woody species and should be considered an important step for pasture restoration.     

Germination and seedling survival in fens undergoing succession

We studied mechanisms of vegetation change in fens subject to succession from open water to floating mats and finally herbaceous rich-fens. Earlier research showed that these systems are characterized by transient seed banks. Our main question was whether seedlings of later successional fen stages are already present in earlier stages, remaining subordinate in the vegetation until conditions become suitable for them. If, however, conditions during succession change in a way that only a limited set of species can survive as seedlings during each of the successional stages, no seedling bank will exist. The transient character of the seeds would then imply that seeds will not germinate and will subsequently die and that seeds that have germinated in the “wrong” stage will not become established. We hypothesized that: (1) germination and seedling survival of fen species are significantly better in the successional fen stage for which these species are characteristic, (2) as a consequence no seedling banks occur in these fens. In a field experiment, seeds of five characteristic fen species in the standing vegetation of three successional fen stages i.e. raft fen, quaking fen and rich fen were sown in each of these stages in a turf pond in the Tienhoven area, The Netherlands. Germination and seedling survival were measured over two growing seasons together with environmental variables. Germination was higher in the “own” stage for all species groups as was survival for quaking fen species and rich fen species. For both these stages, percentage of germination and survival of four out of five characteristic species were significantly higher in the “own” stage. Germination and survival can be considered stage-dependent and it was concluded that seedling banks do not exist in these fens. Site-specific environmental variables act as a sieve and differentiate on species presence already during early life history stages. We found clues that the environmental sieve acts at the level of nutrient availability, tolerance for high sulphide concentration and light climate. Because of the transient seed bank and absence of a seedling bank in these fen wetlands, successful establishment of species necessitates a continuous dispersal of characteristic species until the environmental conditions permit establishment. This also implies that species of the whole successional sere should be present within dispersal distance.     

Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

Environmental controls of raised-bog vegetation in the Baltic boreo-nemoral zone

Raised peat bogs harbor unique vegetation types in specific hydrological conditions. Environmental controls of peat bog vegetation are relatively well known for the boreal zone, while in the European boreo-nemoral zone healthy raised bogs are nowadays very rare. By contrast, Latvia, located in the transition zone between the nemoral and the boreal biomes, still has a large number of active raised bogs. The aim of the present study was to characterize the environmental controls on raised bog vegetation structure, species composition and ecology in Latvia. The study includes 17 raised bogs, where vascular plants, bryophytes and lichens were recorded in 480 sample plots and related to environmental variables (microtopography, litter cover, electric conductivity, pH, and macroelements Na, K, Ca, Mg and P in bog surface waters). The factor best explaining total species richness and composition was microtopography, which also affected most other explanatory factors. Thereby total species richness and cover were highest on hummocks. However, the importance and direction of the effects of microtopography and the other factors differed between vegetation groups. When disregarding microtopography, species composition was most strongly correlated with alkaline ions and litter cover and for bryophytes also with vascular plant cover. The present study is the first wide-scale study in Latvia relating raised bog vegetation to environmental conditions.     

Rodent-favored cache sites do not favor seedling establishment of shade-intolerant wild apricot (Prunus armeniaca Linn.) in northern China

Seed dispersal by scatter hoarders is widely assumed to involve directed dispersal, in which microhabitats selected for caching also benefit seedling establishment and growth. However, in many systems, this may not be true if animal-favored cache sites do not match the safe sites for plants, or if cache sites benefit the plant in one life stage and not another. Here, we investigated whether cache sites selected by rodents are favorable for seedling establishment and growth of shade-intolerant wild apricot (Prunus armeniaca Linn.) in northern China. We tracked tagged seeds and compared the germination and growth of seedlings from rodent-cached seeds with that of naturally established seedlings in a secondary forest and shrubland stand. Rodents preferred to cache seeds under shrubs with medium canopy cover (31–60 %) in litter substrate in the secondary forest, and under shrubs with high canopy cover (>60 %) in soil or litter substrate in the shrubland stand, neither of which conveyed an advantage for seedling establishment. Although fewer caches were made along shrub edges, or under low canopy cover (≤30 %) in the secondary forest, or along shrub edges, open areas of grass, or under low canopy cover in the shrubland stand, these cache sites consistently contributed to higher survival rates. The microhabitats of grass, soil, or low canopy cover significantly promoted the emergence, survival, and growth of naturally established seedlings. Our results are best explained by the conflicting demands of rodents for caching seeds in more secure sites and P. armeniaca’s high-light requirements for seedling recruitment. We argue that the relationship between favored cache sites for seed hoarders and safe sites for plants will often not match but may still allow a reasonable rate of establishment and regeneration.     

森林凋落物对林地微生境的影响

评述了凋落物改变林地微生境的效应．凋落物减弱了林地地表及表层土壤(2mm厚)中的光照,改变了光谱特性,降低了PPFD和Pfr．缩小了日温差,提高了土壤湿度和肥力,增加了毒素和土壤动物的活动等,但凋落物改变生境的效应强度与凋落物组成、厚度、分解状态和周围生境(如植被、气候、地形和地貌等特性)等有关,在自然森林生态系统中,凋落物对林地微生境的效应因凋落物、植被、气候、地形和地貌的多样化变得更为复杂,而凋落物对植被天然更新的影响也因其对生境改变效应的复杂性而具有多样性和综合性,最后,对凋落物改变生境效应的复杂性和有关研究进行了展望。     

A vegetation switch as the cause of a forest/mire ecotone in New Zealand

Abstract. Vegetation switches are those processes in which there is positive-feedback between vegetation and environment, i.e. a vegetation state modifies its environment producing conditions more favourable to itself (Wilson & Agnew 1992). Switches can produce and maintain abrupt ecotones between plant communities. Such a sharp ecotone exists between beech-podocarp forest and mire vegetation, both on deep peat, in southwest New Zealand. One such site was examined. There was no apparent explanation for the ecotone in the present topography nor in the substrate. Levelled transects through the forest demonstrated that most seedling establishment occurred on dead fallen tree boles. These microsites were significantly richer in N, P and K than the wet sump microsites. We argue that this is a mechanism whereby trees can become established in the forest, but not in the open mire. In the forest, the presence of trees ensures the presence of dead-log microsites on the ground, permitting tree seedlings to grow. In the mire, there are no such micro-sites, and trees cannot establish. The ecotone may be sharpened because of the presence of an ecotonal band of the small tree Leptospermum sco-parium between forest and mire. This species can reproduce vegetatively by root suckers in the mire. Its boles are light, and even if they fall to the mire surface they are not thick enough to form a substrate for tree-seedling establishment. The larger tree species may be prevented from falling onto the mire by the wind-sheltering of the forest, and by the zone of Leptospermum. The postulated process would represent a new kind of water-/nutrient-mediated switch, of Type 1 (‘One-sided’). It may occur in many waterlogged forests worldwide.     

Key variables controlling the vegetation of a cool-temperate mire in northern Japan

Abstract. In the cool-temperate Bibi Mire, Hokkaido, Japan, valley fens and flood-plain fens have quite different vegetation. The main variables controlling the vegetation were all hydrological: mean water level, water level fluctuation and surface water flow. Chemical factors such as electrical conductivity, dissolved oxygen and related peat decomposition were less important. The pH was about neutral and has little effect. The flood-plain fen developed under fluctuating water table conditions. The dominant species are Calamagrostis langsdotffii and Carex pseudocuraica. When temporal inundation occurs in the rainy or typhoon seasons, the submergence stimulates bud germination of the stoloniferous C. pseudocuraica, which can rapidly elongate its stolons upward. Some large floating peat mats occurred in the flood-plain fen zone. On these mats some Alnus japonica saplings establish and patches of alder forest can arise. Here the water level was higher than in the peripheral alder forest zone. The valley fen is dominated by Carex lasiocarpa var. occultans and/or C. limosa. It is formed under stable water table conditions in the inundated parts of the mire -where the non-inundated wet areas are dominated by alder trees. In the area where the surface water is flowing, these two fen sedges grow in deeper water since the high oxygen content is considered to compensate the flooding stress.