Kai Jensen

Effects of light competition and litter on the performance of Viola palustris and on species composition and diversity of an abandoned fen meadow

Theeffects of light competition and litter on seedling recruitment and theperformance of established individuals were examined in Violapalustris. This polycarpic perennial plant was a common component ofspecies-rich fen meadows in northwestern Germany until the middle of thiscentury, but today is considered to be regionally endangered. From summer 1996until summer 1998 a bi-factorial field experiment combining three standing croptreatments (mowing, thinning, control) with two litter layer treatments (litterremoval, no litter removal) was carried out in an abandoned fen meadow toinvestigate the effects of these factors both on V.palustris and on aboveground species composition and diversity.MANOVAs (multivariate analysis of variance) revealed significant treatmenteffects for the performance of V. palustris related toexperimental manipulation. Mowing increased vital attributes including thenumber of rosette leaves, the percentage of individuals with chasmogamous andcleistogamous flowers and the number of rhizomes. In contrast, the length of theleafstalk, mean leaf area and the length of rhizomes were negatively associatedwith mowing. Litter removal significantly increased the number of V.palustris seedlings. The removal of the litter layer already resultedin an increase in aboveground species richness in the first year of theexperiment. After three years of experimental manipulation, both abovegroundspecies richness and diversity were positively associated with mowing.Redundancy Analysis showed that typical wet meadow species (Cardaminepratensis, Lychnis flos-cuculi, Lotuspedunculatus) and species of mesotrophic fens (Violapalustris, Agrostis canina, Potentillapalustris) were positively influenced by mowing. It can be concludedthat V. palustris recruitment and growth is promoted by aregular disturbance regime on fen meadows. The conservation of many typicalspecies of wet meadows and fens in northwestern Germany depends on moderategrazing or cutting. These management types prevent successional changes whichlead to an increase in standing crop and the development of a litter layer, bothof which can cause local extinction.

Consequences of abandonment for a regional fen flora and mechanisms of successional change

Until the 1960s, species-rich vegetation on minero- trophic peaty soils (fen sites) were characteristic of the alluvial plains in Schleswig-Holstein (Northwest Germany). Today, many of these habitats undergo successional changes due to abandonment. Vegetation development after abandonment can be characterized as a sequence of different successional stages and described in terms of a successional model. Successional stage I includes grazed, mown and recently abandoned sites without dominants. Stages II and III are characterized by the dominance of highly competitive herbaceous species while stage IV consists of woody vegetation. Ca. 3000 phytosociological relev6s were assigned to the respective successional stages. Mean cover values were calcu- lated for 250 species of the regional fen flora and assigned to successional categories according to their changes in cover in the successional series. According to our results 141 species decrease during succession, while 100 species were restricted to early successional stages and 85 species increased. Aban- donment of all fen sites in Schleswig-Holstein will probably lead to the regional loss of 23 species of the fen flora. To identify mechanisms underlying successional change, the successional categories were correlated with life history traits and ecological requirements of the species. Results indicate that both light competition and limitation of sexual reproduction of small-seeded species might play a major role in the decrease and extinction of species during succession. Finally, conservation strategies for endangered species in a cultural landscape are discussed.

Water-borne seed transport and seed deposition during flooding in a small river-valley in Northern Germany

Summary Water-borne seed transport and seed deposition during flooding were studied in the Upper Eider river (N-Germany) by direct sampling of the rivers seed content with aquatic seed traps and by analysing the number of deposited seeds on sedimentation mats which were exposed near the river on the soil surface during a flooding period of approx. three weeks. The number of seeds which were transported at the surface of the river Eider was continuously analysed by four aquatic seed traps for a period of 20 weeks (July–December 2000). To test the capture rate of these traps, a recapture experiment with colour marked seeds of Helianthus annuus L. was carried out. During the investigation period approx. 9000 seeds of 76 species were captured by the four aquatic seed traps. The number of trapped seeds varied both spatially (across the river profile) and temporally. Considering this variation and the capture rate of the traps, the water-borne seed transport was estimated to be 3139 seeds per week and meter of the river profile. The seed deposition during a flood in early spring 2002 was analysed by using 20 sedimentation mats. To distinguish effects of seed dispersal into patches from outside from seed rearrangement within patches, the water-borne seed transport was excluded from one half of the mats by fencing them with a woven fabric which was permeable for water but not for floating seeds. Outside of the exclosures 152 viable seeds of 26 species were deposited on the sedimentation mats while only one single seedling was found on mats from which water-borne seed transport was excluded. The results demonstrate that hydrochorous dispersal processes might play an important role in connecting otherwise fragmented populations in periodically flooded habitats along rivers.

DORMANCY PATTERNS, GERMINATION ECOLOGY, AND SEED-BANK TYPES OF TWENTY TEMPERATE FEN GRASSLAND SPECIES

The germination ecology, including primary and secondary dormancy, mortality of seeds, and seed-bank type, of 20 fen grassland species from Northern Germany was investigated using a combination of burial and germination experiments. To analyze primary dormancy and effects of after-ripening (dry storage for 28 days) on freshly matured seeds, germination was measured at two fluctuating temperature regimes (15/25°C and 5/15°C) in the light and in darkness. Temporal changes in dormancy were investigated by burying seeds at 8–10 cm depth in nylon bags, exhuming samples at bimonthly intervals over a period of two years, and analyzing germination at the above-mentioned temperature and light treamtents. Additional seed samples were retrieved five years after burial to determine seed longeity and seed-bank type. Freshly matured seeds of all species except Bromus racemosus showed primary dormancy. Dry after-ripening significantly increased germination in Parnassia palustris and Triglochin palustre. Mortality of buried seeds of Bromus racemosus, Sanguisorba officinalis, and Succisa pratensis reached 100% within 12 months (transient seed banks). All other species germinated both in the first and in the second growth period after burial and (with the exception of Briza media) showed annual changes in dormancy. For Silene flos-cuculi and Juncus filiformis, dormancy cycles were detected only in the dark treatments. Most species had a lower percentage germination in darkness than in the light, and the greatest suppression of germination in darkness was found in the small-seeded species (Juncus filiformis and Parnassia palustris). The retrieval of seed samples after five years revealed that most of the fen grassland species examined have short-term, persistent seed banks and thus are buffered against years of poor seed production and/or seedling survival. In addition, a large proportion of the species maintain long-term, persistent seed banks from which re-establishment is possible if management practices and site conditions of degenerated fen grasslands become appropriate following restoration measures.

Seed dispersal in fens

ABSTRACT Question: How does seed dispersal reduce fen isolation and contribute to biodiversity? Location: European and North American fens. Methods: This paper reviews the literature on seed dispersal to fens. Results: Landscape fragmentation may reduce dispersal opportunities thereby isolating fens and reducing genetic exchange. Species in fragmented wetlands may have lower reproductive success, which can lead to biodiversity loss. While fens may have always been relatively isolated from each other, they have become increasingly fragmented in modern times within agricultural and urban landscapes in both Europe and North America. Dispersal by water, animals and wind has been hampered by changes related to development in landscapes surrounding fens. Because the seeds of certain species are long-lived in the seed bank, frequent episodes of dispersal are not always necessary to maintain the biodiversity of fens. However, of particular concern to restoration is that some dominant species, such as the tussock sedge Carex stricta, may not disperse readily between fens. Conclusions: Knowledge of seed dispersal can be used to maintain and restore the biodiversity of fens in fragmented landscapes. Given that development has fragmented landscapes and that this situation is not likely to change, the dispersal of seeds might be enhanced by moving hay or cattle from fens to damaged sites, or by reestablishing lost hydrological connections. Nomenclature: Anon. (2004).

effects of pollination and pollen source on the Seed Set of Pedicularis palustris

Habitat alteration can deteriorate plant-pollinator interactions and thereby increase the risk of population extinction. As part of a larger study on the effects of changes in land use on fen grassland vegetation, factors influencing the seed set of a short-lived, endangered wetland plant,Pedicularis palustris, were studied. We conducted field pollination experiments in one large and one small population. To investigate the effect of pollen source on seed set, individual flowers of caged plants were left unpollinated or were pollinated with pollen from the same flower, the same population or another population. To study pollen limitation and flower display, whole plants were subjected to pollinator exclosure, hand pollination or natural pollination. Self-compatibility was high, but differed between populations (61% and 97% of seed set after cross-pollination within populations). Cross-pollination between populations did not significantly alter seed number per capsule. Pollinator exclosure resulted in a very low seed set (<15% of natural seed set), despite high self-compatibility. The most likely explanations for high self-compatibility in combination with low autofertility are geitonogamy as reproductive assurance, selective neutrality of self-compatibility and phylogenetic constraints. Because of low autofertility, the seed set inP. palustris depends on pollinators. In the study populations, natural pollination was clearly sufficient for maximum seed production per plant, but seed set per capsule was significantly pollen-limited in the smaller population. Plants in this population also had a higher maximum percentage of simultaneously open flowers than those of the large population (31% vs. 13%), while flower longevity was generally extended without pollination. It is concluded thatP. palustris may influence pollinator behaviour and therefore the risk of pollen limitation by flower display.P. palustris showed a flexible reaction to differing pollination regimes without losses in overall seed set in the study populations.

Effects of litter removal and mowing on germination and establishment of two fen-grassland species along a productivity gradient

To study the factors that limit the occurrence of species in fen grasslands, the effects of mowing and litter removal on germination, growth and establishment of two common species (Silene flos-cuculi andLotus pedunculatus) were analyzed along a productivity gradient at seven sites in Northern Germany. The sites differed in nutrient availability, vegetation composition and standing crop. In autumn 2002 a field experiment was set up at each site, including factorial combinations of two treatments (litter removal, mowing). Seeds and juvenile individuals of both species were transferred within the treatment combinations. The number of germinated and transplanted individuals was recorded during the 2003 and 2004 seasons and biomass of survived plants weighed in late summer 2004.Mowing and litter removal had positive effects on the number of germinated seeds of both species. The effect of litter removal was mostly significant at sites with a comparatively low nutrient availability and standing crop (350 g/m2), while the effect of mowing was more important at the site with the highest standing crop (1000 g/m2). In some cases the presence of litter had a positive effect on the survival of individuals, which may be caused by extremely dry weather conditions in 2003.The experiment showed that plant species reaction on mowing and litter removal differed both according to their life stage and the site conditions. These differences could be interpreted in context with Shifting Limitation Hypothesis because the abolition of seed limitation by sowing and creation of additional gaps for germination was most efficient at low productive sites, while the high standing crop reduced first of all the establishment probability, which could be increased by mowing.

Plants mediate soil organic matter decomposition in response to sea level rise.

Tidal marshes have a large capacity for producing and storing organic matter, making their role in the global carbon budget disproportionate to land area. Most of the organic matter stored in these systems is in soils where it contributes 2-5 times more to surface accretion than an equal mass of minerals. Soil organic matter (SOM) sequestration is the primary process by which tidal marshes become perched high in the tidal frame, decreasing their vulnerability to accelerated relative sea level rise (RSLR). Plant growth responses to RSLR are well understood and represented in century-scale forecast models of soil surface elevation change. We understand far less about the response of SOM decomposition to accelerated RSLR. Here we quantified the effects of flooding depth and duration on SOM decomposition by exposing planted and unplanted field-based mesocosms to experimentally manipulated relative sea level over two consecutive growing seasons. SOM decomposition was quantified as CO2 efflux, with plant- and SOM-derived CO2 separated via δ(13) CO2 . Despite the dominant paradigm that decomposition rates are inversely related to flooding, SOM decomposition in the absence of plants was not sensitive to flooding depth and duration. The presence of plants had a dramatic effect on SOM decomposition, increasing SOM-derived CO2 flux by up to 267% and 125% (in 2012 and 2013, respectively) compared to unplanted controls in the two growing seasons. Furthermore, plant stimulation of SOM decomposition was strongly and positively related to plant biomass and in particular aboveground biomass. We conclude that SOM decomposition rates are not directly driven by relative sea level and its effect on oxygen diffusion through soil, but indirectly by plant responses to relative sea level. If this result applies more generally to tidal wetlands, it has important implications for models of SOM accumulation and surface elevation change in response to accelerated RSLR.

Patterns of wetland plant diversity along estuarine stress gradients of the Elbe (Germany) and Connecticut (USA) Rivers

Background: Estuaries are characterised by salinity gradients and regular flooding events. These environmental factors form stress gradients, along which species composition changes. Aims: Analyse and compare patterns of plant species diversity along the estuarine salinity and flooding gradients of the Elbe and Connecticut Rivers. Methods: Vegetation was sampled at three elevations (low, mid, high) in five sites of each marsh type (fresh, brackish, salt) in both estuaries. Patterns of species density (SD) and evenness (E) along the gradients were analysed and compared between the two estuaries with three-factor ANOVAs. Results: The regional species pool was 33% higher for the Connecticut than for the Elbe. SD of fresh marshes (19 ± 2.2) was more than twice in the Connecticut than in the Elbe. We found an overall increase in SD from low to high elevation and from salt to freshwater marshes in both estuaries. However, SD and E were strongly depressed at intermediate elevations in the Elbe fresh and brackish marshes. Conclusions: Although diversity patterns in the two estuaries show overall similarities, patterns of SD and E differ, when particular elevational zones and marsh types are compared. We hypothesise this to be due to evolutionary and historical influences on the regional species pools, shaping the impact of local biotic and abiotic processes.

Fen Management and Research Perspectives: An Overview

A fen has vegetation that is actively forming peat and is fed by ground-or sur- face water (Joosten and Clarke 2002). In Europe a “fen meadow” is a ground- or surface water-fed mown grassland that does not form peat, since it was formed after modest drainage of a fen or it developed on a predominantly moist soil (Grootjans and Van Diggelen 1995). Therefore, fens and fen mead- ows are considered to be different ecosystems by most European authors. Others do not make a distinction between fens and fen meadows because the species composition of both ecosystems may overlap considerably (Wheeler et al.1995). In North America, fens dominated by tussock-forming sedges are referred to as ‘sedge meadows’, which are often grazed. Since there is not yet scientific agreement on whether sedge meadows are fens or fen meadows, we will refer to them as fens in this chapter.