Late holocene history and vegetation dynamics of a floodplain alder carr: A case study from eastern Bohemia, Czech Republic

Vegetation development in the lowland floodplain alder carr “Na bahně” (eastern Bohemia, the Czech Republic) has been studied by means of pollen and macrofossil analyses and combined with vegetation analysis performed over the last 70 years. Local successional changes started with an oxbow lake (160 cal BC) which has later terrestrialised (630 cal AD). Then it changed from a typical alluvial fen into aSphagnum-dominated spring mire (950 cal AD) supplied by water arising from a river terrace surrounding the locality from three sites. In the centre of this wetland a small patch of alder carr developed (100 cal. AD), showing some tendency towards cyclic succession. The alder carr alternated several times with an openCarex fen (1100 cal AD to recent). The last fen-to-alder carr transition has been documented by direct observation during this century. Possible autogenic and allogenic factors driving the succession are discussed. The model of autogenic cyclic succession corresponds well with direct field observations and can be used to interpret alder carr structure, its dynamics, and function.     

How soil characteristics and water quality influence the biogeochemical response to flooding in riverine wetlands

Although phosphate concentrations have been reduced, the rivers Meuse and Rhine are still polluted with sulphate, which most probably affects vegetation development in newly created riverine wetlands. The influence of flooding with river water rich in sulphate was tested on three soil types from floodplains of the river Meuse using flow-through and batch experiments. Soils were selected for contrasting concentrations of iron and organic matter and originated from a floating fen (iron-poor, organic), an alder carr (iron-rich, organic) and a clay pit (iron-rich, low in organic matter). Flooding induced mobilisation of phosphate. Sulphate only enhanced this effect in the alder carr soil, where sulphide and phosphate competed for binding to iron. Only in the floating fen soil did the addition of sulphate result in the formation of free sulphide, which reduced the growth of Glyceria maxima, serving as a phytometer. In addition, the floating soil started to sink, due to falling methane concentrations. In the different soil types methane production was hampered by the presence of more favourable electron acceptors such as sulphate in the water and Fe(III) in the soil. It was concluded that the effects of inundation with sulphate-polluted water strongly depend on the soil type: under iron-poor circumstances, free sulphide may accumulate, leading to phytotoxicity, while in soils rich in iron, sulphide toxicity is prevented, but phosphate availability may be increased. In addition, shortage of easily degradable organic matter can limit the formation of potential toxicants such as ammonium, iron and sulphide. Results are discussed in terms of their implications for nature management.     

PLEISTOCENE HISTORY OF THE BRITISH VERTEBRATE FAUNA

• 1 This review covers the Pleistocene history of British non-marine Pisces, Amphibia, Reptilia and especially Mammalia, which alone have a good fossil record. Aves are also briefly discussed.
• 2 The fossil material available is often inadequate for purposes of taxonomy and identification. Further problems arise because many groups of Mammalia have undergone rapid evolution during the Pleistocene.
• 3 In this paper the fossil record is related to the currently accepted stratigraphic table of the British Pleistocene (Shotton & West, 1969). Wherever possible, fossil records have been assigned to pollen assemblage zones. Throughout, emphasis is placed on the relationship between faunal history and vegetational history, as determined from fossil pollen and macroscopic plant remains.
• 4 Although fossils are relatively scarce in the fluviatile and lacustrine deposits of open sites, compared with the often rich cave assemblages, the stratigraphy of the former is usually much clearer and the sediments commonly contain pollen. It is difficult to correlate cave sequences with those of open sites.
• 5 It is important to take into account possible bias in a fossil assemblage according to its mode of accumulation, e.g. assemblages from occupation sites may represent only those animals which were hunted by man.
• 6 Lower Pleistocene vertebrates are rather poorly-known. The majority of fossils are from the marine Crags of East Anglia (Pre-Ludhamian to Pastonian) and a single cave assemblage of this age is known (Dove Holes). Few records can be related to particular stages, but a few finds from Easton Bavents are assigned to Antian and Baventian stages.
• 7 Early Middle Pleistocene vertebrates are represented mainly by the rich assemblages from the marine and fresh-water Weybourne Crag and Cromer Forest Bed Series (Baventian to Early Anglian) of Norfolk and Suffolk. The East Runton fauna appears to be of pre-Cromerian (?Pastonian) age. A good fauna is known from the type Cromerian deposits at West Runton (zone Cr 11). A few records are available for zone ?Cr III and one for the Early Anglian. The assemblages from other localities appear to represent more than one stage at each site, e.g. the so-called ‘Bacton Forest Bed’ fauna is composite, including both Cromerian and ?Pastonian taxa. Outside East Anglia one open site (Sugworth) and one cave fauna (Westbury) of probable Cromerian age are known.
• 8 Many of the fossils found in lacustrine and river-terrace deposits of the Middle and Upper Pleistocene glacial-interglacial succession (Anglian to present day) can be assigned to particular stages, zones or even subzones. Cave assemblages rarely predate the Ipswichian. No pre-Devensian records are available for either Scotland or Ireland. The Anglian fauna is very poorly known. The Hoxnian is represented principally by the Clacton (zone Ho IIb) and Swanscombe faunas. The Baker's Hole deposit, the basal gravels of the Summertown-Radley Terrace and the Glutton and Bear Strata in Tornewton Cave have yielded faunas of probable Wolstonian age. The early Ipswichian is poorly represented (Selsey), many fossils are known from zone Ip Iib (e.g. Trafalgar square, Swanton Morley, Aveley), there are a few records from early zone Ip III (Aveley, Swanton Morley) and fairly good faunas from zone Ip III/IV (Histon Road, Stutton). Several open and cave-site faunas resemble those of zones Ip II and Ip III and the assemblages from Ilford, Brundon, etc., appear to date from the end of this interglacial. The Early Devensian is represented by the Wretton fauna and probably by some cave faunas. Middle Devensian faunas are fairly well known (e.g. Upton Warren) and there are a number of records for the Late-Devensian (Ballybetagh, High Furlong, Nazeing). Many cave faunas date from the Middle or Late Devensian. Good faunas are available from the early Flandrian, zone F1 I (e.g. Star Carr). The present-day native fauna (zone F1 111) is also discussed.
• 9 The main faunal characteristics for each subdivision of the Pleistocene are summarized in the Conclusions. There is a major faunal change between the predominantly Tertiary fauna of the Red Crag Nodule Bed (Probably Pre-Ludhamian and older) and that of the Red Crag (Pre-Ludhamian and Ludhamian). There appears to have been comparatively little change in fauna through the rest of the Lower Pleistocene but the more intense climatic fluctuations of the Middle and Upper Pleistocene were accompanied by rapid faunal change and the appearance of characteristic ‘steppe-tundra’ faunas in the Wolstonian and Devensian cold stages. The Late-Devensian and Flandrian faunas are impoverished in comparison to earlier stages. This may be partly due to the activities of man as well as climatic and vegetational changes.
• 10 There is usually good agreement between fauna and vegetational conditions when these can be compared, but a few taxa (e.g. Cricetus cricetus, Equus) have clearly changed their ecological requirements during the Pleistocene. Changes of fauna in response to vegetational changes within interglacials are known from the Hoxnian and especially the Ipswichian. The ‘steppe-tundra’ vegetation of cold stages was accompanied by a mixture of animals nowadays extinct or living in either steppe or tundra.

     

Palaeoenvironmental research on diatoms in early and middle Holocene deposits in central North Holland (The Netherlands)

In the geological research of the Holocene coastal deposits of The Netherlands, diatoms are used to reconstruct the sedimentary facies, palaeo-tide levels and salinity gradients during the deposition of the sediments. In this paper, the results of diatom research from 4 borings, taken from the early and middle Holocene deposits of central North Holland are presented. The oldest marine influenced sediments in the area are the deposits of the ‘Velsen layer’, a clay layer rich in organic matter and deposited about 8000–7000¹⁴C years before present (BP) at a depth of 20–12 m below present mean sea-level. This clay layer was formed in a shallow, permanently submerged environment with a limited tidal influence (lagoonal or pond-like conditions). The salinity changed from brackish/freshwater to marine/brackish. The younger sandy and clayey sediments, formed about 7000–4500 BP at a depth of 16–3 m below present mean sea-level, are classified as ‘tidal channel’ and ‘interchannel’ deposits. It is argued, on the base of both diatom and non-diatom criteria, that the lower and middle parts of the interchannel deposits in the central area of the palaeo-tidal basin of North Holland were formed in a subtidal environment. The upper part of the interchannel deposits and the deposits at the fringe of the North Holland tidal basin were formed in the intertidal zone or, at the fringe of the basin, even in the supratidal zone. The salinity in the North Holland tidal basin during the sedimentation of the tidal channel and interchannel deposits was marine/brackish to marine. This study indicates that diatoms, besides their palaeoecological applications, have chronostratigraphical significance (on a regional scale). The diatomsCymatosira belgica andActinoptychus splendens appear to be a useful (eco)stratigraphical marker in the Holocene coastal deposits of The Netherlands and Belgium.     

Short-Term Vegetation Dynamics of Alnus Dominated Peatlands: a High Resolution Palaeoecological Case Study from Western Pomerania (NE Germany)

Actual ecological research postulates for alder carrs a cyclic alternation of Alnus tree vegetation with open fen or Salix dominated vegetation. Such cycles are also indicated in palaeo-ecological studies, but normally the temporal resolution of these studies is insufficient to resolve the duration of short-term cycles in vegetation development. This paper presents a high resolution palaeoecological study (including pollen, macrofossils and non-pollen palynomorphs) of a Late Holocene wood peat section from the small, long-term Alnus dominated peatland ‘Heger Soll’ in the ‘Rodder Forst’ in Western Pomerania (NE Germany) to reconstruct short-term vegetation changes. During a time-span of ca. 800 years, sedge-dominated fen vegetation types alternated with two phases of Alnus carr and one phase of Salix shrubland. The alder carr decline coincided with the beginning of intensified human activity in the surroundings of the mire and was probably connected to increased water discharge resulting from large-scale deforestation, after which willow scrub and sedge fen became established. Growth of Alnus trees was associated with prolonged phases of reduced human impact and probably less water supply. This study shows that human impact on the uplands surrounding the mire and on the alder carr itself may explain the observed “cyclic” vegetation development of alder carrs, willow scrubs and sedge fens in Central Europe.     

Quantitative studies of modern wet-ground molluscan faunas from Bossington, Hampshire

There have been few quantitative studies of modern Mollusca in Britain, especially from wet-ground areas. This is particularly true of the English chalklands, which is surprising given the focus of sub-fossil molluscan research in that area. Those studies that are available highlight the difficulty of separating molluscan faunas from structurally complex habitats such as fen and carr. Quantitative molluscan data from a variety of wet-ground habitats situated within the chalklands of southern England were analysed using an indirect ordination technique, Principal Components Analysis. This demonstrated that the separation of faunas from structurally complex wet-ground habitats is possible. The numerical separation does not rely on indicator species, but is a function of variation of the predominant species. It is suggested that further studies of this type might aid the interpretation of sub-fossil Mollusca from wet-ground contexts.     

Fossil blackflies from Pleistocene interglacial deposits in Norfolk, England (Diptera: Simuliidae)

ABSTRACT. The occurrence is reported of fragmentary simuliids among Pleistocene fossil arthropod assemblages recovered from deposits in Norfolk of Ipswichian (last) interglacial age, 130 000–120 000 years BP. The condition of each of the nineteen available specimens is described, and sixteen of them are photographically illustrated. Specimens are identified as Simulium s. l. but reliable identification beyond generic level is precluded by lack of diagnostic taxonomic characters. The palaeoenviron-mental significance of simuliids in Quaternary fossil assemblages is briefly discussed; it is thought that the simuliids indicate a palaeoclimate no colder, and possibly warmer, than today.     

The interpretation of fen carr pollen diagrams: pollen–vegetation relationships within the fen carr

Sediments deposited in fen carr environments are frequently used in pollen-based reconstructions of vegetation history, although few studies of modern pollen–vegetation relationships in fen carr have been undertaken to aid the interpretation of fossil assemblages. It is often assumed that the wetland pollen signal will reflect the overall composition of the fen carr system although, given the closed canopy context, it may actually be dominated by local vegetation elements. This paper seeks to determine whether the vegetation heterogeneity within a modern fen carr is reflected in the pollen signal of the wetland taxa. Vegetation composition was recorded around a series of moss polster sample points at Calthorpe and Wheatfen, two areas of fen carr in the Norfolk Broadland, UK. The spatial interrelationship between the wetland components of the vegetation and their palynological equivalents in the moss polster pollen assemblages is explored through trend surfaces and a correlation based statistical test (the Mantel Test). The representation of the major vegetation components in the pollen record is highly variable. Values for Alnus glutinosa, a dominant canopy species, range from 5.3% to 73% of total land pollen (TLP) at Calthorpe and 2.5–61% TLP at Wheatfen. In part, this variation reflects the enhanced input of pollen from dry land sources close to the wetland edge although the variation recorded at points equidistant from the dry land margin suggests that other processes are also influential. The strongest positive associations in the Mantel Test are for under-storey and ground flora elements that are poorly represented in the pollen record. Taxa for which there is no association include those with few vegetation occurrences and those for which a large proportion of the pollen appears to have been derived from outside the study areas. We conclude that modern pollen samples collected from fen carr broadly reflect the vegetation composition of the study areas. However, the input of pollen from fen carr species is spatially inconsistent. Differences between the sample points can be attributed to the distribution and pollination biology of the under-storey and ground flora elements and the structure of the vegetation around sites. The results caution against assuming that changes in the pollen representation of wetland taxa, in fossil sequences constructed from fen carr deposits, necessarily reflect changes in the overall composition of the community.     

Shingle-based wetlands at Dungeness

WATERS, S. J. P. & FERRY, B. W., 1989. Shingle-based wetlands at Dungeness . At Dungeness, a unique development of open water and natural wetland has occurred on the shingle beaches. Over 700–1000 years, low-lying areas have changed from saline lagoons into freshwater lakes which are now largely infilled by reedswamp and sallow carr. Since about 1970, the water-table has been lowered, by a combination of water abstraction and hydrological changes associated with gravel extraction. Water-table lowering has accelerated colonization by reedswamp and sallow carr, and has caused loss of sedge-rich marshland. Possible methods are suggested for promoting development of such marshland in artificial excavations.     

Reed die-back related to increased sulfide concentration in a coastal mire in eastern Hokkaido,Japan

A drastic decline of Phragmites australis was observed along the middle reaches of Ichibangawa River in Kiritappu Mire, eastern Hokkaido, Japan, during the last 50 years. In an area of ~30 ha, reed-sedge vegetation and alder forest have been replaced by bare soil and patches of salt marsh vegetation. A gradual increase in frequency of flooding by brackish water probably was the ultimate cause of the vegetation change. We measured redox potentials and oxygen and sulfide concentrations in soil profiles using needle electrodes. Measurements were carried out in areas where reed has disappeared and in sites where reed stands were still healthy. The concentration of selected ions in the surface water was also measured at various sites. Surface water in low-lying areas was clearly influenced by seawater. Very high sulfide concentrations were measured in bare peat sites (more than 600 mol l^–1), which exceeded P. australis tolerance 2 – 3 times. In a healthy reed zone adjacent to an area with poor fen vegetation, sulfide concentration in the rooting zone of Phragmites was also high (300–400 mol l^–1), particularly during the night. The fact that Phragmites in this zone was still healthy indicates that sulfide did not reach toxic levels in the direct vicinity of the roots. Sulfide that is produced in this area is probably fixed by iron, which is supplied through a continuous discharge of iron-rich groundwater. An increase in frequency of flooding by brackish water could be related to ongoing subsidence of this part of the Pacific coast which is located at the Kuril subduction zone. Sea level rise could also contribute to a stronger inflow of seawater into the mire system.     

Tropical forest and mangrove history from southeastern Mexico: a 5000 yr pollen record and implications for sea level rise

A 250 cm long core from El Palmar, a swamp area located along the Rio Hondo river in the south of the Yucatan Peninsula, near the Belizean border, reveals the environmental history of the mangrove and tropical forest of the last 5000 years. The period between 5000 and 4600 b.p. shows sandy deposits, which form the early infill and development of the swamp. A medium-statured tropical forest covered the area and members of the Moraceae and Fabacaeae dominated this early forest. The period between 4600 and 4000 b.p. presents a clear change to a mangrove system with Conocarpus erecta and Rhizophora mangle as dominant trees. This vegetational change is due to flooding of the Rio Hondo river, which deposits sediments of high salinity due to higher sea-level. The medium-statured forest became established at some distance from the swamp area. After 4400 b.p. C. erecta appears as the dominant mangrove species and the R. mangle stands are less predominant in the area. The tropical forest was close to the swamp area and was mainly composed of members of the Moraceae, Arecaceae and Fabaceae as dominant taxa of this vegetational mosaic.     

Long‐term effects of drainage and initial effects of hydrological restoration on rich fen vegetation

Questions: What vegetational changes does a boreal rich fen (alkaline fen) undergo during a time period of 24 years after drainage? How is plant species richness affected, and what are the changes in composition of ecological groups of species? Is it possible to recover parts of the original flora by rewetting the rich fen? Which are the initial vegetation changes in the flora after rewetting? What are the major challenges for restoration of rich fen flora after rewetting? Location: Eastern central Sweden, southern boreal vegetational zone. Previously rich fen site, drained for forestry purposes during 1978–1979. The site was hydrologically restored (rewetted) in 2002. Method: Annual vegetation survey in permanent plots during a period of 28 years. Results: There were three successional stages in the vegetational changes. In the first stage there was a rapid (< 5 years) loss of rich fen bryophytes. The second step was an increase of sedges and early successional bryophytes, which was followed by an increase of a few emerging dominants, such as Molinia caerulea, Betula pubescens and Sphagnum spp. After rewetting, there are indications of vegetation recovery, albeit at slow rates. Depending on, for instance, initial species composition different routes of vegetation change were observed in the flora after drainage, although after 24 years, species composition became more homogenous and dominated by a few species with high cover. Conclusion: Major changes have occurred after changes in the hydrology (drainage and rewetting) with a severe impact on the biodiversity among vascular plants and bryophytes. Several rich fen bryophytes respond quickly to the changes in water level (in contrast to vascular plants). The recovery after rewetting towards the original rich fen vegetation is slow, as delayed by substrate degradation, dispersal limitation and presence of dominant species.     

Long-term vegetation dynamics and the infilling process of a former lake (Švarcenberk, Czech Republic)

Natural lakes are a rare phenomena within extraglacial areas of Central Europe. Almost all have been completely terrestrialized during the Holocene. This paper deals with one such former lake, located in southern Bohemia. Its extensive lacustrine and peat deposits were subjected to a multidisciplinary study that resulted in high-resolution pollen, macrofossil, algal and sediment-chemistry data interpreted in terms of past climate, geomorphology, soil, and regional vegetation development over the last 16,000 years. Against the background of these large-scale processes, local development took place, comprising the lake’s ontogeny from an arctic-type ecosystem hosting pioneer aquatic communities, through a highly diversified mosaic of eutrophic hydrosere habitats (shallow pools,Phragmites andCarex fen, alder carr), towards an oligotrophic mire that started to dome over the terrestrialized lake. At every individual development stage, specific processes characterized ecosystem function and composition: during the Late-Glacial with its rapid climatic changes, external forces induced the major stresses; while during the Holocene, autogenic changes of the wetland ecosystem played the most important role.     

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.     

Palaeoecological evidences from foraminifers and ostracods on Late Quaternary sea-level changes in the Ombrone river plain (central Tyrrhenian coast,Italy)

The multidisciplinary study of borehole Ombrone S1, sampled in the Grosseto plain, near the Ombrone river mouth, provides new data about sea-level changes during the Holocene in central Tyrrhenian Sea. The palaeontological analysis of the borehole highlights data concerning the foraminifers and ostracods useful to the palaeoenvironmental interpretation. Moreover, geochemical analyses (δ¹³C and δ¹⁸O) were performed on Ammonia parkinsoniana tests and Cyprideis torosa valves. Integrating the results of the palaeontological, lithological and geochemical data, it is possible to single out the environments related to the transgressive trend. Borehole Ombrone S1 is characterised by basal deposits of continental settings, which includes some brackish water marsh, may be related to a slight sea-level oscillation, followed by fluvial gravel ones testifying the deltaic progradation of the Palaeo-Ombrone river. Above, two distinct trangressive events, which were divided by a phase of still-stand, took place. The first one, above the gravel, was responsible for the establishment of the brackish water conditions due to the presence of a barrier-lagoon system, which was finally filled up by terrigenous input. The second one produced, at first, lagoon settings near the sea and, successively, deltaic-related settings for the submersion of the barrier. The littoral sandy bar at 10 m of core-depth represents the first sure evidence of the alluvial plain progradation, which develops up to the core-top.     

The first pollen diagram from the Hoher Fläming, Brandenburg (Germany)

A 72 cm peat core from a spring mire reveals 3300 years of vegetation history in the Hoher Fläming, a landscape formed by the penultimate glaciation in southeastern Brandenburg. Primeval beech-oak forest dominated from a.d. 550 to 1200, prior to local forest clearance. Existing maps showing the natural and potential natural vegetation as pine and oak need to be revised. Local woodland cutting in the 12th century and grassland use since the 15th century affected the development of the helokrene (spring-fed) mire deposits from alder carr to mesotrophic fen vegetation and then to the present-day Sphagnum fallax-Juncus acutiflorus wet meadows. Vegetation and mire history link together the Hoher Fläming and adjacent landscapes on a northwest-southeast gradient of suboceanic to subcontinental climate. Moreover it represents a geographical and altitudinal transition of lowland and hill vegetation from 30 to 200 m a.s.l. to the montane belts of the palynologically well investigated Harz mountains. However, regarding some trees and herbs, the Hoher Fläming does not belong to the Hercynian region and type of vegetation.     

Postglacial evolution of vegetation and environment in the Scheldt Basin (northern Belgium)

The infill of abandoned river channels is the main information source on postglacial vegetation and landscape evolution in inland northern Belgium. Palynological studies of these deposits also contribute to a better understanding of the environmental context of the numerous archaeological sites in this area. The results are presented of palynological analyses, including pollen, spores, non-pollen palynomorphs and microcharcoal from the palaeochannel infill at seven sites in the Scheldt and Durme valleys in northern Belgium. Based on the regional vegetation evolution and radiocarbon dating, a biozonation for the Scheldt Basin is constructed encompassing the final Late-glacial and Holocene period. The local taxa show a hydroseral succession from open-water environment, through reed or sedge fen to carr forest. Salix and Viburnum opulus were important elements in the Early Holocene riparian and carr vegetation, but are often underrepresented in pollen diagrams from larger valleys. There are indications for human disturbance and agriculture between 7,000 and 6,400 cal bp. The simultaneous decline of elm and lime in the study area around 4,000 cal bp can be linked to human impact.     

Microenvironmental conditions for Japanese alder seedling establishment in a hummocky fen

In this article, we focused on hummocky microtopography as a prominent feature of mires and explored the microenvironmental conditions suitable for alder seedling establishment. Japanese alder (Alnus japonica) forest is widely distributed in wetlands in northern Japan. However, because alder seedlings are rare in mires, alder population dynamics and conditions that favor the establishment of alder seedlings are still unknown. The study was conducted in northern Japan at a site in mesotrophic mire. We surveyed the seedling density, the microenvironmental conditions (light, litter cover, and soil quality), and the density of dispersed seeds in alder forest and in the adjacent herbaceous fen. In addition, we performed a laboratory experiment to examine the germination characteristics of alder. Seedlings grew only on hummocks in alder forest. The percentage of litter cover on hummocks was lower than in hollows, and the density of dispersed seeds in alder forest was much higher than in herbaceous fen. Seeds of Japanese alder germinated under both light and dark conditions, and the germination rate were high under light and high-temperature conditions. Our results suggest that litter cover may inhibit seedling establishment and hummocks that characterized by less litter cover are suitable place for the establishment of seedlings. We conclude that hummocky microtopography and abundant seed rain in the mire enable the establishment of Japanese alder seedlings.     

Species richness and succession of aquatic beetles in some kettle-hole ponds in northern Sweden

Ten closely-located kettle-hole ponds in northern Sweden were ordered along a vegetational, successional gradient from the present-day representation of the six vegetational units distinguished. Different rates of vegetational development caused by differences in depths of basins have made possible the long-term existence of early successional stages. A fact that has offered the possibility of studying most of the vegetational succession of these habitats.
The number of water beetle species inhabiting each pond were correlated to successional stage and area of pond. Mean species numbers decreased from 34 to 14 per pond with increasing maturity of the vegetational succession. This decrease in species richness of water beetles is caused by a decrease in number of habitats or vegetational units present. The increased floral conformity is mainly due to a more stable water-level caused by lower rates of bottom seepage from the accumulation of organic deposits. Besides the extended zonation of littoral vegetation, a high water-amplitude is also connected with recurrent nutrient pulses caused by the faster detritus composition during drought periods. Thus the changing water-levels contribute to a high water beetle diversity through the maintenance of both a high habitat complexity and a high production.
A positive correlation between pond size and number of water beetle species were only found if the fen ponds were excluded. From this it is concluded that the area effect is here a good predictor of species richness only within ponds of a similar successional stage.