Changes along a disturbance gradient in the density and composition of propagule banks in floodplain aquatic habitats

This study used germination methods to examine the density, species composition and functional composition of propagule banks in a series of riverine wetland aquatic habitats subject to varying degrees of hydrological and management-related disturbance. Under permanent inundation (the conditions prevailing at most sites during the growing season) propagule germination and species richness was low, with floodplain perennials and helophytes particularly affected. Densities of floodplain annuals were largely maintained through continued germination of a few flooding tolerant species. On damp mud (conditions associated with hydrological instability) total seedling number and species richness increased significantly, but species richness of germinating hydrophytes declined. Mean seedling density at 0–0.1m depth was 15450 ± 4400 m–2, reaching a maximum (162 050 m–2) in temporary backwaters. Annual (e.g., Lindernia dubia, Cyperus fuscus) and facultative ruderal species (e.g., Lythrum salicaria and Alisma plantago-aquatica) predominated. Vertical zonation of the propagule bank was weakly developed. The numbers of individuals and species germinating varied significantly between sites. The seasonal, most intensely disturbed sites (temporary backwaters) supported a numerically large, species-rich propagule bank based on floodplain annuals, while the permanent, less disturbed sites (ditches and an oxbow pond) had a small, species-poor propagule bank composed of hydrophytes and helophytes supplemented by allochthonous seed inputs. Sites intermediate on the gradient had a propagule bank dominated by facultative amphibious, ruderal hydrophytes. The composition of the seed bank and the established vegetation was most similar at the heavily disturbed sites where the seed bank was maintained by vigorously fruiting annuals and supplemented by inputs from temporary habitats upstream. At permanent sites much of the propagule bank composition could be accounted for by inputs of floodborne seed from the immediately adjacent floodplain. The established vegetation at such sites appeared to be maintained mainly by vegetative propagation with recruitment from the propagule bank likely only after severe disturbance. The potential contribution of functionally diverse propagule banks to sucessional processes within fluvially dynamic floodplain aquatic habitats is emphasised.     

Impact of flooding on potential and realised grassland species richness

In order to reduce flood risk, river management policies advise floodplain restoration and the recreation of water retention areas. These measures may also offer opportunities for the restoration of species-rich floodplain habitats through rewetting and the restoration of flood dynamics. The potential to enhance biodiversity in such flood restoration areas is, however, still subject to debate. In this paper we investigate whether flooding along a small altered lowland river can contribute to the potential and realised species richness of semi-natural meadows. We compare the seed bank and vegetation composition of flooded and non-flooded semi-natural meadows and test the hypothesis that flooding contributes to an input of diaspores into the meadow seed banks, thereby promoting seed density and potential species richness. Furthermore we hypothesise that, where habitat conditions are suitable, flooding leads to a higher realised species richness. Results showed that seed densities in flooded meadows were significantly higher than in non-flooded meadows. The seed banks of flooded meadows also contained a higher proportion of exclusively hydrochorous species. However, the seed bank species richness, as well as the species richness realised in the vegetation did not differ significantly between flooded and non-flooded meadows. Finally, the seed bank and standing vegetation of flooded sites showed larger differences in species composition and Ellenberg nitrogen distribution than non-flooded sites. From these results we conclude that, although flooding does contribute to the density and composition of the seed bank, most imported seeds belong to only a few species. Therefore, it is unlikely that flooding substantially enhances the potential species richness. Furthermore, even if new species are imported as seeds into the seed bank, it seems unlikely that they would be able to establish in the standing vegetation. However, it is unclear which factors impede the establishment of imported species in the vegetation. The implications of our findings for flood meadow restoration are discussed.     

Comparison of soil seed banks of habitats distributed along an altitudinal gradient in northern Iran

In this study we investigated the variations in soil seed banks along an altitudinal gradient in the Alborz mountains, Iran, covering three habitats from lower to upper altitudes: forest, forest-subalpine grassland ecotone and subalpine meadow. In each habitat from 1850 to 2400 m, 20 quadrats were established along four transects, and the above-ground vegetation and the germinable seed banks were determined. Results show that the similarity between seed bank and vegetation was lowest in the ecotone located at intermediate altitudes. Together with the contrasting highest density and species diversity of seeds at these altitudes, the ecotonal role of this habitat was confirmed.We found evidence that lower altitudes could act as storage for seeds of some species growing at higher altitudes; the role of the ecotone was more prominent as a reserve for the meadow plant seeds than the role of the forest as a reserve for seeds of the meadow and ecotone habitats. Soil seed banks, particularly from the ecotone, can be used for restoring vegetation in some degraded sites.     

Plant diversity in a calcareous wooded meadow – The significance of management continuity

Questions: What is the contribution of management continuity during the last 30–40 years to variation in species diversity and composition of a calcareous wooded meadow plant community? Is tree cover related to species diversity and composition of the herbaceous layer? What are the effects of local soil gradients on species diversity? Location: Laelatu calcareous wooded meadow, Western Estonian coastal zone. Methods: Plant community composition was assessed in 150 1 m × 1 m plots, located at 30 sites with known management history within Laelatu meadow (7 ha). Light and soil conditions and relative altitude were measured at each plot. DCA was used to analyse variation in species composition and general linear mixed models to analyse the effects of management and environmental parameters on diversity. Results: Management continuity was the primary determinant of plant community composition, followed by light conditions and soil parameters. Species richness, diversity and evenness are positively dependent on management continuity. Spatial autocorrelation is important as well. Diversity started to decline under the tree canopy where 50% or less irradiation reached the level of the herbaceous layer. We did not find significant effects of soil conditions on small‐scale diversity. Conclusions: Management continuity, together with the cover of the tree layer, are the most important determinants of diversity. Despite grassland stands with different management history are located side by side, the regeneration of diversity and composition of plant communities after restoring regular management practices is a slow process.     

Digging deep for diversity: riparian seed bank abundance and species richness in relation to burial depth

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Convergence of community composition during secondary succession on Zokor rodent mounds on the Tibetan Plateau

Aims The community succession theory is much debated in ecology. We studied succession on Zokor rodent mounds on the Tibetan Plateau to address several fundamental questions, among them: (i) During secondary succession, does the community composition converge towards one community state or multiple states depending on the initial colonization? (ii) Do mound communities located in different background communities exhibit different assembly trajectories?Methods In a sub-alpine meadow, we investigated a total of 80 mound communities at several successional stages in three different background communities resulting from different management histories and compared their changes in species composition. The distribution of plant communities over time was analyzed with quantitative classification and ordination methods. The co-occurrence patterns of species were evaluated at each successional stage, and the degree of convergence/divergence among communities was obtained by calculating two beta-diversity indices.Important findings During secondary succession, species richness of mound communities changed over time, and this change was dependent on the background community. Five life-form groups exhibited different dynamic patterns in species richness and plant cover. Community composition and the degree of species co-occurrence between communities increased over time since disturbance. There was much variation in species composition at earlier stages of succession, but communities on older mounds became more similar to each other and to their surrounding vegetation over the course of secondary succession. Post-disturbance succession of Zokor mound communities transitioned from 'multiple alternative states' to 'background-based deterministic community assembly' over time. Tradeoffs between competition and colonization, as well as the characteristics of different life-forms and mass effects within a limited species pool are the mechanisms responsible for convergence of mound communities.     

Order and disorder in the river continuum: the contribution of continuity and connectivity to floodplain meadow biodiversity

Aim Aspects of connectivity and continuity operating in the River Meuse were analysed for their contribution to the biodiversity of the floodplain. From this analysis of the diversity and composition of the meadow communities, we aimed to derive effective biodiversity conservation strategies. Location The River Meuse is one of the larger rivers in the European Western Plains ecoregion. The alluvial plains of the river have a long history of cultivation, and for these plains the floodplain meadow vegetation is a highly appreciated and valuable nature conservation asset. Method We sampled floodplain meadows from 400 km of the six geomorphic reaches of the middle to lower course of the River Meuse. For each, 50 vascular plant relevés were recorded, representing the spectrum of floodplain meadow communities of that reach. Beta diversity was calculated to quantify similarity in species pools between the reaches. A dissimilarity formula was used to determine the turnover between the reaches, and these dissimilarities were compared with a Mantel test to detect whether species composition of the floodplain meadows exhibited connectivity and continuity between the reaches. Species richness for the floodplain vegetation data of the reaches was compared with data for riparian invertebrate communities. The vegetation data sets were ordinated using detrended correspondence analysis (DCA) to reveal patterns in the floodplain meadow species composition, and the DCA axes were related to plant functional groups and population strategies. The axis scores of the species and plots were linked to river and plant species traits. Results We did not observe an overall continuity trend in similarity, nor one in diversity in a downstream direction. Lateral connectivity was highlighted by the dissimilarity between the reaches and in the influxes of species from adjacent ecoregions. The DCA ordination showed statistically significant separations between reaches and between the plant functional groups. The second DCA axis related to the longitudinal gradient of the river, whereas the first axis showed stronger correlations with river and plant species traits. We termed this axis the ‘disorder axis’. Plant invaders and avoiders are located at the left side of the disorder axis, whereas the true river‐adapted categories of resisters and endurers are at the right extremity. Contributions to the disorder were identified in terms of connectivity with adjacent ecoregions and the physical disturbance regime of natural and anthropogenic perturbations, resulting in community changes between the reaches. Main conclusions We concluded that a single overall strategy for biodiversity conservation of the river or its floodplains is not feasible. Strategies can, however, be derived for separate river reaches based on functional groups in the communities, the disorder characteristics of the reach, and the influence of surrounding ecoregions.     

Plant functional trait responses to grassland succession over 25 years

Abstract. Many grasslands and arable fields have been abandoned in Europe in the last decades. So far, however, ecologists have not been able to establish one general, unifying theory for successionally induced changes in species composition following abandonment. In fact, the course of succession seems to be unique for each site and year. Here we focused on the analysis of plant functional traits in order to detect general trends in trait responses to succession which would prove independent of site characteristics. We studied permanent plot series of 14 grassland sites in SW Germany that had been abandoned for 25 yr. Species composition as well as the course of succession varied significantly among the grasslands. Response to succession was analysed for the following traits: ‘plant height’, ‘canopy structure’, ‘specific leaf area’ (SLA), ‘storage organ’, ‘vegetative spread’, ‘plant persistence’, ‘seed bank longevity’, ‘start of flowering’, ‘duration of flowering’ and ‘seed mass’. We compared a univariate with a multivariate approach. In the univariate approach, attributes of each trait were analysed separately employing GLM whereas in the multivariate approach all traits were handled together in NMS. Both analyses indicated similar trait responses to succession. There was a significant increase in the proportion of species characterized by the following attributes: plant height > 0.6 m, leaves distributed regularly along the stem, vegetative spread > 100 mm, start of flowering later than May, duration of flowering 1–2 months and seeds of either low or high mass.     

The impact of flooding regime on the soil seed bank of flood‐meadows

Abstract. We assessed the significance of flooding for the floristic composition of seed banks in flood‐meadows of the northern valley of the Upper Rhine. We compared three hydrological compartments of the alluvial plain, consisting of the regularly flooded land between the river and low summer dykes (functional flood‐plain), the occasionally flooded land between summer dykes and high winter dykes (hybrid floodplain) and the land behind the winter dykes, which is now only submerged by ascending groundwater (fossil flood‐plain). Due to their different flooding regime, the three compartments should differ with respect to the prevailing conditions of diaspore input. The seed density of soil samples increased with the duration of flooding in the three compartments, while species richness and the proportion of species not occurring in the vegetation was constant. The increase in seed density can be largely attributed to an increase of disturbance indicators, which are present in the above‐ground vegetation and capable of forming a long‐term persistent seed bank. No effects of flooding on the composition of seed banks in the three flood‐plain compartments were found. The differences in seed bank composition can be largely explained by corresponding differences in above‐ground vegetation and former and present‐day meadow management. Seeds of species absent from above‐ground vegetation can be attributed to the local species pool present in the immediate vicinity of the study plots. We discuss consequences of the results for the restoration of species‐rich flood‐plain meadows.     

Impacts of Land Abandonment on Vegetation: Successional Pathways in European Habitats

Changes in traditional agricultural systems in Europe in recent decades have led to widespread abandonment and colonization of various habitats by shrubs and trees. We combined several vegetation databases to test whether patterns of changes in plant diversity after land abandonment in different habitats followed similar pathways. The impacts of land abandonment and subsequent woody colonization on vegetation composition and plant traits were studied in five semi-natural open habitats and two arable habitats in six regions of Europe. For each habitat, vegetation surveys were carried out in different stages of succession using either permanent or non-permanent plots. Consecutive stages of succession were defined on a physiognomic basis from initial open stages to late woody stages. Changes in vegetation composition, species richness, numbers of species on Red Lists, plant strategy types, Ellenberg indicator values of the vegetation, Grime CSR strategy types and seven ecological traits were assessed for each stage of the successional pathway. Abandonment of agro-pastoral land-use and subsequent woody colonization were associated with changes in floristic composition. Plant richness varied according to the different habitats and stages of succession, but semi-natural habitats differed from arable fields in several ecological traits and vegetation responses. Nevertheless, succession occurred along broadly predictable pathways. Vegetation in abandoned arable fields was characterized by a decreasing importance of R-strategists, annuals, seed plants with overwintering green leaves, insect-pollinated plants with hemi-rosette morphology and plants thriving in nutrient-rich conditions, but an increase in species considered as endangered according to the Red Lists. Conversely, changes in plant traits with succession within the initially-open semi-natural habitats showed an increase in plants thriving in nutrient-rich conditions, stress-tolerant plants and plants with sexual and vegetative reproduction, but a sharp decrease in protected species. In conclusion, our study showed a set of similarities in responses of the vegetation in plant traits after land abandonment, but we also highlighted differences between arable fields and semi-natural habitats, emphasizing the importance of land-use legacy.     

Seed rain of fleshy and dry propagules in different habitats in the temperate rainforests of Chiloé Island,Chile

In temperate rainforests on Chiloé Island in southern Chile (42°S), most canopy trees bear fleshy, avian‐dispersed propagules, whereas emergent tree species have dry, wind‐borne propagules. In the present study, the following hypothesis was tested: regardless of species, fleshy propagules are deposited in greater numbers in canopy gaps and in forest margins and hence have a more heterogeneous seed shadow than wind‐dispersed propagules. To test this hypothesis, the seed rains of these two types of propagules were compared in the following forest habitats: (i) tree‐fall gaps (edges and centre); (ii) forest margins with adjacent pastures; and (iii) under closed canopy (forest interior). Seed collectors (30‐cm diameter) were placed in two (15 and 100 ha) remnant forest patches (n = 60–100 seed collectors per patch) distributed in the four habitats. Seeds were retrieved monthly from each collector during two reproductive seasons (1996, 1997). In both years, the seed rain was numerically dominated by two species with dry propagules (Laureliopsis philippiana and Nothofagus nitida) and three species with fleshy fruits (Drimys winteri, Amomyrtus luma, and Amomyrtus meli). The seed shadows of the two species with dry, wind‐dispersed seeds differed markedly. Seeds of L. philippiana were deposited predominantly in canopy openings, whereas N. nitida seeds fell almost entirely in the forest interior. The fleshy‐fruited species, Drimys and Amomyrtus spp., had similar seed deposition patterns in the various habitats studied, but the between‐year differences in seed rain were greater in Drimys winteri than in Amomyrtus spp. Although no more than 10% of fleshy‐fruited propagules reached the margins of the patch, approximately 7% of these were carried there by birds. Every year, canopy gaps (pooling data from edges and centres) concentrated approximately 60% of the total seed rain of both propagule types in both forest patches. Forest margins received less than 20% of the total seed rain, which was largely dominated by fleshy‐fruited species. Seed shadows were a species‐specific attribute rather than a trait associated with propagule type and dispersal mode.     

Role of substrate and landscape context in early succession: An experimental approach

Both local site conditions and landscape context influence the course of succession, but there is a lack of experimental studies on the relative importance of these two factors. It is hypothesised that convergence vs. divergence in succession is determined by the interplay of site factors, such as type of substrate and the nature of the surrounding landscape. In order to evaluate the role of substrate and surrounding landscape in the initial development of vegetation, experimental plots with tertiary clay, sand, peat, sterilised local soil and undisturbed local soil as a control were established in two contrasting regions, and the cover of all the species present was recorded annually for 10 years. In early succession, vegetation was affected by both the substrate and surrounding landscape, but their effects resulted in different trends. The importance of the substrate gradually decreased, while that of the landscape context increased. In the course of succession the vegetation between the two regions diverged and converged within each region. We concluded with regard to the divergence vs. convergence dichotomy in succession: if contrasting habitats occur in the same or similar landscapes, convergence is expected, whereas if similar or the same habitats are located in contrasting landscapes, divergence is expected. For the remaining combinations, i.e. contrasting habitats in contrasting landscapes or the same habitats in the same or a similar landscape, successions may exhibit no or only slight divergence or convergence.     

Seed dispersal in red deer (Cervus elaphus L.) dung and its potential importance for vegetation dynamics in subalpine grasslands

Free-ranging large herbivores can influence vegetation dynamics through seed dispersal within and among habitats. We investigated the content of germinable seeds in the dung (endozoochory) of red deer (Cervus elaphus L.), the most ubiquitous wild ungulate throughout the European Alps, and compared the results with the species composition of the vegetation type in which the dung was dropped. The study was conducted in the subalpine zone of the Swiss National Park and included the three most important vegetation types for red deer: (i) intensively grazed short-grass vegetation, (ii) less intensively grazed tall-grass vegetation, and (iii) adjacent conifer forest understory vegetation. Seeds of 47 species, mostly from small-seeded herbaceous species, were recorded in dung samples with three species accounting for 65% of germinated seeds. Our results confirmed the hypotheses that (H1) small-seeded species were more likely to occur in red deer dung than larger-seeded species, though seed size was unrelated to seed density, (H2) red deer dung contained mostly seeds from short-grass vegetation, with seed species composition in dung collected from any vegetation type being most similar to species composition of relevés from short-grass vegetation, and (H3) seeds were less likely to be dispersed between vegetation types than within vegetation types, with dung dropped in short-grass vegetation having a different species composition and containing over twice as many seeds as dung dropped in the other two vegetation types. These results collectively support the hypothesis that red deer endozoochory contributes to maintaining short-grass vegetation, the favoured grazing sites of hinds in the Swiss National Park, by increasing propagule pressure of seeds from herbaceous forage species adapted to endozoochory relative to other species and especially those from later stages of secondary succession.     

Vegetation responses to environmental conditions in floodplain grasslands: Prerequisites for preserving plant species diversity

We studied plant species responses to environmental conditions in floodplain grasslands (alliance Cnidion) of the river Elbe (northern Germany) in order to describe relationships between species composition/diversity and underlying site gradients. We analysed a total area of 639 ha of floodplain grasslands, using 98 sampling plots. Vegetation responses to environmental conditions were examined by means of DCA. The relationships between species diversity and environmental conditions were examined using regression analyses. Our results show that species composition and species richness are mainly affected by present-day utilization, by the phosphate supply and by the productivity of stands. Species richness decreases significantly under grazing, with increasing phosphate supply and productivity of stands. Intermediate productivity levels, mowing and high inundation frequencies promote floodplain target species. N_inorg availability and base availability have only a minor impact on species composition/species diversity. Productivity is closely related to the amount of plant available phosphate. Low phosphate concentrations in the topsoil are particularly characteristic of Cnidium dubium floodplain meadows. Therefore, it is likely that in floodplain grasslands with low to intermediate productivity, phosphate is the most important factor influencing plant competition and plant species diversity. We hypothesize that an increased phosphate supply increases stand productivity and, hence, the competition for light, thus excluding weak competitors. This interpretation is supported by the present-day distribution of C. dubium floodplain meadows of the river Elbe: these are concentrated in the hybrid (between summer and winter dykes) and fossil floodplain (landward side of winter dykes), but have been replaced in the functional floodplain by grass-dominated plant communities as a result of a steady increase in the river water phosphate content. It is, therefore, only in the hybrid and fossil floodplain grasslands that species diversity may be preserved in the long term, as the P input is lower here than in the functional floodplain.     

Dispersal,diversity and distribution patterns in pioneer vegetation: The role of river‐floodplain connectivity

Question: Are there changes in dispersal patterns in floodplain pioneer vegetation with effects on seedling number, species richness and species composition along a gradient of declining river‐floodplain connectivity? Location: Middle Elbe river floodplain, Germany. Methods: An experiment with five treatments was set up along a gradient of declining river‐floodplain connectivity, partitioning seedlings into three groups: (1) emerging solely from water dispersed seeds, (2) from wind/animal dispersed seeds and (3) from the soil diaspore bank. Two controls were established: without any manipulation and exclusion of all seeds. The results were compared with those of vegetation and soil sampling to evaluate the representativeness of the experimental sites in terms of species composition, diversity, seedling number and soil parameters. Results: Water dispersal and the soil diaspore bank were the major dispersal strategies shaping floodplain pioneer vegetation at the Middle Elbe river. The number of seedlings, species richness and the variation in species composition in these habitats depend on the degree of connectivity. The seedling number and species richness is highest in sites of permanent or almost permanent exchange with the main channel, where water dispersal additionally contributes to the number of seedlings grown from the soil seed bank. Conclusion: The results underline the importance of river‐floodplain ecotones as sink habitats for water‐dispersed seeds. Considering the strongly reduced river‐floodplain interactions due to dykes and other engineering structures, management strategies are necessary to improve connectivity and the renewal of fluvial land forms.     

How does the propagule bank contribute to cyclic vegetation change in a lakeshore marsh with seasonal drawdown?

Lakeshore marshes around Liangzi Lake, in the middle reach of the Yangtze River, China, experience annual changes of water level of c. 1.5 m. During the drawdown period, the vegetation is structured by helophytes and emergents; during the rainy season when the dams are closed (June–September) the marshes are flooded and their vegetation rapidly changes to be come dominated by submerged, floating-leaved and tall emergent species. The species composition and abundance of both the marsh seed bank and the vegetative propagule bank were compared with those of the drawdown and flooded vegetation types. These data provided a test of the predictive power of van der Valk's model of northern temperate seasonal vegetation change in a subtropical, freshwater wetland with cyclic vegetational change. The abundant species were detected in the propagule bank. The seed bank was found to determine the species richness of both types of the vegetation, whereas the vegetative propagule bank consisted of the dominants of the drawdown vegetation. Water depth conditions, and the composition of seed and vegetative propagules banks together determine the structure of the standing vegetation during drawdown and flooding. van der Valk's succession model was found to predict the seasonal vegetation change reasonably well. The Chi-square test showed no significant difference between predicted vegetation and actual vegetation in both drawdown and flooding periods.     

Propagule pressure and environmental conditions interact to determine establishment success of an invasive plant species,glossy buckthorn (<Emphasis Type="Italic">Frangula alnus</Emphasis>), across five different wetland habitat types

Many invasive plant species are able to establish within a wide range of community types. This establishment success depends on high propagule pressure and successful recruitment of seedlings following propagule dispersal into receptive environments. This study aimed to investigate interactions between propagule pressure and environmental resistance to seedling recruitment of the invasive shrub, glossy buckthorn (Frangula alnus Mill.), over a range of wetland habitat types. We measured propagule deposition using seed traps and recruitment success using sown plots, while characterizing vegetation and abiotic environmental conditions in five adjacent wetland habitat types. Drier habitats, which included Cedar Swamp, Shrub Carr, and String, had lower resistance to buckthorn establishment than the wetter Flark and Cattail Marsh. The drier habitats supported more woody species and provided more raised hummock surfaces essential for successful buckthorn recruitment and establishment. Propagule pressure was also higher in dry habitats that supported higher densities of adult glossy buckthorn, while long-distance dispersal into areas with low adult density was uncommon. Natural recruitment was highest in sites with intense propagule pressure, but experimental sowing of seeds demonstrated that buckthorn establishes in wet sites with higher resistance if propagule pressure is increased and seeds are deposited on hummocks. This study demonstrates the affinity of glossy buckthorn for drier wetland sites, and provides empirical evidence that environmental resistance can be overcome by higher propagule pressure.     

Invasive sweetclover (<Emphasis Type="Italic">Melilotus alba</Emphasis>) impacts native seedling recruitment along floodplains of interior Alaska

Sweetclover (Melilotus alba) is a non-native legume that has formed dense and extensive patches along several rivers in Alaska. Our research objective was to determine if sweetclover impacts recruitment of native seedlings in floodplain habitats. To determine if sweetclover impacted recruitment, we conducted a removal experiment along two rivers in interior Alaska. When compared to areas where sweetclover was removed, areas with sweetclover had approximately 50% greater mortality of native seedlings, 25% less recruiting species, and a significant reduction in the quantity of light available to seedlings on floodplain surfaces. To determine if sweetclover shading was a mechanism that limited seedling recruitment, we grew eight common early-successional floodplain plant species in a greenhouse under a range of lighting conditions that were representative of shading under sweetclover. We observed that species restricted to the earliest seral stages of floodplain succession experienced greater reductions in biomass than species that persist into later stages of floodplain succession. Shading seedlings in a greenhouse did not lead to mortality during the growing season. However, when seedlings were over-wintered, we observed that greater shading during the growing season can result in higher seedling mortality. Our study indicates sweetclover invasions have created a novel shade environment in early seral floodplain plant communities, which has the potential to alter community composition. To preserve biodiversity and structure of plant communities, sweetclover should be actively managed to prevent its spread onto additional glacial rivers in Alaska.     

Reintroduction of a Rare Plant (Gladiolus imbricatus) Population to a River Floodplain—How Important is Meadow Management?

In seminatural grasslands, the success of reintroduction of locally extinct rare plant species may depend on the ambient management regime. We aimed to study to what extent the success of the restoration of a rare species (Gladiolus imbricatus) depends on management conditions. A management experiment with traditional cutting by scythe and hay removal, mowing (machine cutting and hay removal), mulching (machine cutting without hay removal), spring burning, and unmanaged control, combined with reintroduction of seeds of Gladiolus, was conducted in an Estonian flooded meadow in which the species had become extinct. Seeds were reintroduced in 2003 in all management treatments and populations monitored until 2006. Mulching, mowing, and traditional management resulted in the greatest establishment, whereas the subsequent mortality was not influenced greatly by management regime. The population started to increase in mulching treatment in the third season due to vegetative growth. The results indicate that the establishment of G. imbricatus is primarily seed limited under current conditions, whereas favorable management significantly enhances establishment in a river floodplain meadow. Successful restoration depends on seed addition and proper grassland management—mowing to a height of approximately 15 cm and mulching.     

Early gap successional pathways in a Fagus-Acer forest preserve: pattern and determinants

Abstract. Many theories of forest succession imply that terrestrial plant community composition within a region tends to converge toward a climax community. That is, given similar climatic and edaphic conditions, succession at different sites within an area will lead to comparable species compositions, a pattern referred to as successional convergence. In this study, we examine changes in plant composition within forest canopy gaps over a 17-yr period to identify potential patterns of successional convergence and to ascertain the factors controlling the successional pathway. To do so, we: (1) sampled 36 forest canopy gaps in Hueston Woods Nature Preserve in 1977, 1981, 1985, 1989 and 1993, (2) evaluated changes in the similarity of gap composition over this period, and (3) examined gap composition in each year as a function of variables describing gap habitat, seed source proximity, and disturbance history. Results indicated an initial pattern of successional divergence, with gaps exhibiting increased dissimilarity over the first 10–12 years of succession. We attribute this initial period of divergence to the effects of differential seed inputs from edge individuals and heterogeneity of available light due to differences in gap size. Recent surveys, however, indicated that gap composition has become more similar as competition within gaps has become more intense. In these samples, gap composition is closely linked to site conditions, including slope, soil conditions, and site exposure. Finally, while these patterns may suggest equilibrium-oriented dynamics, non-equilibrium processes such as repeat disturbances are also evident at Hueston Woods and will likely play an important role in determining future successional pathways.