Radial oxygen loss, a plastic property of dune slack plant species

Mean and environmentally induced differences in radial oxygen loss (ROL) were investigated for three pioneer species (i.e. Schoenus nigricans L., Juncus articulatus L. and Samolus valerandi L.) and two late-successional dune slack species (i.e. Calamagrostis epigejos L. and Carex flacca Schreber). These species were grown in a factorial design at conditions differing in moisture, light, nutrient availability and Fe²⁺ and Mn²⁺ concentrations. On average, ROL was twice as low for late-successional species compared to pioneer species, but it increased on average by a factor 1.5 for these species upon flooding while ROL was unaffected by flooding in pioneer species. Other effects of treatments on pioneer and late-successional species groups were insignificant. Species-specific plasticity in ROL was stronger than that of the species groups and varied highly with the different combinations of – interactions between – environmental cultivation conditions flooding, concentrations of Fe²⁺ and Mn²⁺, nutrient and light levels, but was not affected by differences in relative growth rates. Moreover, ROL activity was almost 20×, and significantly, higher for seedlings than for adults of Schoenus nigricans. This implies that ROL of a species depends on the stage of the life cycle and on growth conditions, something that should be considered when determining ROL. ROL activity of most species increased at (a combination of) high nutrient and at low light levels: ROL activity was strongly negatively correlated to the root/shoot ratio, which was presumably caused by a higher gas transport capacity compared to root oxygen consumption at low root/shoot ratios. The plasticity in ROL leads to a highly dynamic rhizosphere in which the oxygen influx is a function of plant species and particularly of the interactions between plant species and environmental conditions.     

[CO2]- and density-dependent competition between grassland species

The predicted ongoing increase of atmospheric carbon dioxide levels is considered to be one of the main threats to biodiversity due to potential changes in biotic interactions. We tested whether effects of intra‐ and interspecific planting density of the calcareous grassland perennials Bromus erectus and Carex flacca change in response to elevated [CO₂] (600 ppm) by using factorial combinations of seven densities (0, 1, 2, 4, 8, 16, 24 tillers per 8 × 8 cm² cell) of both species in plots with and without CO₂ enrichment. Although aboveground biomass of C. flacca was increased by 54% under elevated [CO₂], the combined aboveground biomass of the whole stand was not significantly increased. C. flacca tended to produce more tillers under elevated [CO₂] while B. erectus produced less tillers. The positive effect of [CO₂] on the number of tillers of C. flacca was strongest at high intraspecific densities. On the other hand, the negative effect of [CO₂] on the number of tillers of B. erectus was not present at intermediate intraspecific planting densities. Seed production of C. flacca was more than doubled under elevated [CO₂], while seed production of B. erectus was not affected. Moreover, the mass per seed of C. flacca was increased by elevated [CO₂] at intermediate interspecific planting densities while the mass per seed of B. erectus was decreased by elevated [CO₂] at high interspecific planting densities. Our results show that the responses of C. flacca and B. erectus to elevated [CO₂] depend in a complex way on initial planting densities of both species. In other words, competition between these two model species is both [CO₂]‐ and density dependent. On average, however, the effects of [CO₂] on the individual species indicate that the composition of calcareous grasslands is likely to change under elevated [CO₂] in favor of C. flacca.     

Preference of wet dune species for waterlogged conditions can be explained by adaptations and specific recruitment requirements

The preference of wetland angiosperms for waterlogged soils has been explained by several hypotheses: (1) wetland species are adapted to waterlogging and sensitive to drought; (2) wetland species are tolerant to drought, but inferior competitors at drier conditions; (3) wetland species have narrow moisture optima for recruitment. We tested these hypotheses by the application of permanent and fluctuating water levels to experimental wet dune plant communities with four species that frequently occur in wet dune slacks (Carex flacca, Molinia caerulea, Samolus valerandi and Schoenus nigricans) and one competitive species of drier conditions (Calamagrostis epigejos). After 3 years, fluctuating water levels had led to lower total biomass production than permanent water levels, indicating that switching from aerated to anoxic soil conditions involved physiological costs. The collective biomass of wetland species was highest at permanently waterlogged conditions with interspecific variation in the biomass optima. At the nutrient-poor growth conditions of this long-term experiment, biomass of Calamagrostis was independent of water level treatment, thus the hypothesized competitive superiority of this species at drier conditions could not explain the biomass responses of the wetland species in this study. Instead, this is evidence for an ecological preference of adult wet dune plants for waterlogged to moist conditions. Recruitment of most wet dune slack species occurred in a narrower range of water levels than adult growth, indicating that recruitment requirements also pose a limitation to the distribution of these wet dune species. Incorporation of recruitment into nature management support models may improve their predictions.     

Pedogenesis in coastal wet dune slacks after sod-cutting in relation to revegetation

The development of vegetation and soil was investigated in wet coastal dune slacks in North Holland, The Netherlands (52°36'N, 4°37'E). Sod cutting in the past has created a time series from 1 to 30 years, with an even older undisturbed site as reference. After sod-cutting Schoenus nigricans, a typical pioneer species of wet dune slacks, established and contributed together with Calamagrostis epigejos to more than 85% of the aboveground biomass. At the control site the biomass was 9.5 t dry matter ha^-1.the humus layer increased to a thickness of 11 cm. The concentration of nitrogen, phosphorus and potassium increased in the humus layer without strong effects on the sandy subsoil, in contrast to the strong decalcification in the subsoil. The velocity of the latter process was a factor of hundred higher than reported from dry dunes in the same area.Changes in nutrient amount and availability affected the element concentration in the two dominant plant species. The nitrogen concentration in shoots of S. nigricans was nearly constant over time, whereas that in shoots of C. epigejos declined by 80%. Due to the increasing biomass the size of the N- and P-pool of the biomass increased with the age of the plots.Processes of vegetation development and pedogenesis are not (yet) affected by decalcification as established by the nut mass of S. nigricans. It is concluded that decalcification is not the key factor for the disappearance of rare species. Due to the dominance and the height structure of the two dominant plant species competition for radiation and lack of bare soil for germination are discussed as main reason for losses in biodiversity.It is advised that mowing may be an effective management tool for hampering the soil formation; but sod-cutting may be necessary once in every decade.     

Vegetation development in dune slacks: the role of persistent seed banks

Abstract. The soil seed bank composition was determined at four sites in the dune slack ‘Koegelwieck’ on the Dutch Wadden Sea island of Terschelling. At three different sites in the slack, where sod-cutting experiments down to the mineral sand had been carried out, the established vegetation and seed bank were assessed after 5, 9 and 39 yr of undisturbed development, respectively. In addition, a fourth site in the slack was investigated, where vegetation development had proceeded for 80 yr since plant colonization of bare soil and where nowadays a vegetation dominated by Calamagrostis epigejos and Salix repens occurs. Together these four sites can be regarded as a chronosequence of dune slack formation. Clear time sequences were detected in the seed bank data. Many late successional species showed a significant increase in the number of seeds during the succession. Some of the early successional basiphilous pioneer species such as Anagallis minima, Centaurium littorale, Littorella uniflora, Radiola linoides and Samolus valerandi, showed either a decrease during the time of succession or an optimum in the vegetation while remaining present in the seed bank in low but detectable numbers. They could, therefore, play a role in re-establishment of the vegetation after sod-cutting. One of the target species, Schoenus nigricans, established within a few years after removal of the sod. However, no seeds of this species have been detected in the soil below either of the successional stages. Based on the species disappearance from the established vegetation and based on the independent data of Thompson et al. (1997) an estimation of seed longevity could be made for several Red List species of wet dune slacks.     

Seed survival and persistence on a calcareous land surface after a 32-year burial

Abstract. Of 34 species of vascular plants recorded in an unimproved calcareous grassland community in 1960 and subsequently buried for 32 yr, just six spp. were recovered as viable seeds from the original land surface, following excavation of an experimental earthwork. Five of the six are known or suspected to form persistent seed banks; the status of Carex flacca as a persistent seed is now established. The exceptionally low density of viable seeds at 35 seeds/m² and the paucity of other plant remains confirm that these calcareous soils have supported rapid decomposition. The complete absence of viable seeds known to be short-lived indicated that the macrofauna has played no significant role in the introduction of viable seeds from the modern-day surface down to depths in excess of 1.6 m, at least not in recent years. Because of the precise dates of burial and excavation, the results offer a rare opportunity to confirm laboratory and other indirect observations of long-term seed persistence following burial in temperate climate calcareous soils.     

Genetic variation and hybridization in SwedishSchoenus (Cyperaceae)

Schoenus ferrugineus andS. nigricans have restricted distributions in Sweden and are almost exclusively confined to calcareous fen habitats. AtS. nigricans sites,S. ferrugineus is usually also present, and hybrids are frequently found. In this report, I used allozymes to estimate the amount of gene flow between the two species, and to compare the partitioning of genetic diversity in each of them. Thirteen loci were analysed at eight different enzyme systems. Seven loci were variable between or within the species. The two species had completely different alleles at two of the seven variable loci, whereas there was overlap at five loci. In all, 22 different alleles were found. Six of these alleles were confined toS. nigricans, and five alleles were confined toS. ferrugineus. Nei's genetic identity was 0.55.—InS. ferrugineus, three loci (23%) were polymorphic, and the average number of alleles per polymorphic locus was 2.0 (each polymorphic locus had two alleles). InS. nigricans, three loci (23%) were polymorphic, and the average number of alleles per polymorphic locus was 2.3.—The proportion of genetic diversity due to variation among sites (G _ST) was fairly similar in the two species, mean over loci = 0.12 inS. ferrugineus and 0.15 inS. nigricans. However, the proportion of genetic diversity due to variation among individuals within sites (G _IS) differed markedly between the two species, mean over loci = 0.54 inS. ferrugineus and 0.17 inS. nigricans. Accordingly, there was a much higher individual heterozygosity inS. nigricans than inS. ferrugineus. — Most hybrids were interpreted as F₁ hybrids. However, a small proportion, 0.5–1.6 %, were F_n hybrids or back-crosses.—On the Swedish mainland, all former occurrences ofS. nigricans are extinct, but viable hybrids are still present at a few sites in southernmost Sweden.     

Re-assessment of phosphorus availability in fens with varying contents of iron and calcium

Aim

To further unravel P availability in mineral-rich fens, and test whether high Fe in the soil would lead to low P availability to the vegetation.

Methods

Mesotrophic fens were selected over gradients in Ca and Fe in central Sweden and the Netherlands, to study characteristics of vegetation, pore water and peat soil, including inorganic and organic forms of P, Fe and Al.

Results

Soil Fe was more important than region or soil Ca, and P availability to the vegetation increased from Fe-poor to Fe-rich fens. Contrary to expectations, precipitation of iron phosphates played a minor role in Fe-rich fens. Fe-rich fens were P-rich for three reasons: (1) high P sorption capacity, (2) relatively weak sorption to Fe-OM complexes and (3) high amounts of sorbed organic P, which probably consists of labile P. Also, nonmycorrhizal wetland plants probably especially take up weakly sorbed (organic) P. However, high P did not lead to high biomass or low plant diversity. Fe-rich fens were limited by other nutrients, and high P may help protect the vegetation against Fe-toxicity.

Conclusions

Fe-poor fens are P-poor, irrespective of Ca, and Fe-rich fens P-rich even under mesotrophic conditions. However, high P itself does not endanger Fe-rich fens.

     

Effects of elevated CO2 and phosphorus addition on productivity and community composition of intact monoliths from calcareous grassland

We investigated the effects of elevated CO₂ (600 μl l⁻¹ vs 350 μl l⁻¹) and phosphorus supply (1 g P m⁻² year⁻¹ vs unfertilized) on intact monoliths from species-rich calcareous grassland in a greenhouse. Aboveground community dry mass remained almost unaffected by elevated CO₂ in the first year (+6%, n.s.), but was significantly stimulated by CO₂ enrichment in year two (+26%, P<0.01). Among functional groups, only graminoids contributed significantly to this increase. The effect of phosphorus alone on community biomass was small in both years and marginally significant only when analyzed with MANOVA (+6% in year one, +9% in year two, 0.1 ≥P > 0.05). Belowground biomass and stubble after two seasons were not different in elevated CO₂ and when P was added. The small initial increase in aboveground community biomass under elevated CO₂ is explained by the fact that some species, in particular Carex flacca, responded very positively right from the beginning, while others, especially the dominant Bromus erectus, responded negatively to CO₂ enrichment. Shifts in community composition towards more responsive species explain the much larger CO₂ response in the second year. These shifts, i.e., a decline in xerophytic elements (B. erectus) and an increase in mesophytic grasses and legumes occurred independently of treatments in all monoliths but were accelerated significantly by elevated CO₂. The difference in average biomass production at elevated compared to ambient CO₂ was higher when P was supplied (at the community level the CO₂ response was enhanced from 20% to 33% when P was added, in graminoids from 17% to 27%, in legumes from 4% to 60%, and in C. flacca from 120% to 298% by year two). Based on observations in this and similar studies, we suggest that interactions between CO₂ concentration, species presence, and nutrient availability will govern community responses to elevated CO₂. Received: 12 July 1997 / Accepted: 28 March 1998     

Classification of schoenus communities in South and Southeast Sweden

Summary The Schoenus phytocoenoses of South and South-eastern Sweden have been sampled by 189 relevés of standard areas (4 m²) and divided into clusters using the TABORD program on two homotoneity levels. Two Schoenus associations and five potential nuclei of associations, mainly Schoenus nigricans phytocoena, have been distinguished.The species have been divided into twelve groups on the basis of similarities in distribution amongst phytocoena and assumed environmental preferences. Acidophilous or indifferent species together with common mire species are characteristics of the Oxycoccus-Schoenus ferrugineus association, while basiphilous and wet-meadow species are typical of the Primula-Schoenus ferrugineus association and the Fissidens-Schoenus nigricans phytocoenon. The Primula-Schoenus ferrugineus association is characterized by species associated with Tofieldietalia and Eriophorion latifolii, while in the Oxycoccus-Schoenus ferrugineus association also Scheuchzerio-Caricetalia and Oxycocco-Sphagnetalia species play an important rôle. Another difference between the two associations is that hemicryptophytes and helomorphes dominate in the Primula-Schoenus ferrugineus association, while in the Oxycoccus-Schoenus ferrugineus association dwarf shrubs, hydrophytes and hydromorphes are rather common.The properties of the homotoneity and diversity indices used are compared and their compatibility is discussed.     

Zonation of plant cover and environmental factors in wetlands of the Sanjiang Plain,northeast China

The zonation of depressional and riparian wetlands in the Sanjiang Plain of northeastern China was studied to describe their vegetation composition and environmental variables. We sampled 108 plots in 6 depression and riparian wetlands. Samples were classified into 4 groups using two‐way indicator species analysis (TWINSPAN). Emergent marsh vegetation was characterized by Carex lasiocarpa and C. pseudocuraica, meadow marsh vegetation by tussock species such as Carex appendiculata and C. meyeriana, wet meadow vegetation by Calamangrostis angustifolia and Carex appendiculata, and shrub meadow vegetation by the shrubs Betula fruticosa, Alnus sibirica and Salix rosmarinifolia and the graminids Carex schmidtii and Calamagrostis angustifolia. CCA ordination showed that water table, organic matter and available N were the major factors explaining the vegetation zonation pattern. Compared with other Northern Hemisphere regions, bog and fen vegetation are completely absent due to climatic conditions unfavorable for peat formation. Out of four vegetation types, only the Carex lasiocarpa community and the C. pseudo‐curaica community have been found in other regions. However, at the species level many species are widespread and some species are vicariant or pseudovicariant to other regions in the world. Our study suggests that topography and hydrology may be the most important determinants of the vegetation pattern in this region.     

Changes in soil and vegetation during dune slack succession

Many rare plant species occur in Dutch wet dune slacks, particularly in the Junco baltici‐Schoenetum nigricantis. For nature management it is important to understand the processes controlling the presence of these basiphilous early successional communities, which is why we investigated vegetation and soil development during succession in coastal dune slacks. We compared 12 chronosequential stages of 0, 2, 4, 9, 10, 13, 25, 30, 43, 60, 70 and 85 yr in five different dune slack systems. In four of these locations turf had earlier been removed in order to restore the basiphilous pioneer stage. The main variation in the vegetation is related to the acidification/soil enrichment gradient and the salinity/maritime gradient. During succession, organic matter accumulates and acidification takes place. Maritime influence can buffer the soil and postpone the succession of basiphilous pioneer communities for many years. A significant correlation with age was found for 18 variables. Multiple regressions predicted changes in the vegetation (dependent variables: biomass, cover of Salix repens, Calamagrostis epigejos and Schoenus nigricans) as a function of acidification, organic matter accumulation, increase in available P and presence of Na in the soil. We conclude that natural ageing of the vegetation and the associated process of accumulation of biomass drive succession in this hydrosere. The underlying soil processes are acidification and organic matter accumulation. During succession dominance shifts from S. nigricans to S. repens or C. epigejos. Maintenance of the pioneer character of the habitat is only possible by local intervention or by natural or man‐induced dune forming. The effect of sod‐stripping depends on the environmental conditions and, in case of acidification, success is limited. Succession can be postponed by mowing.     

Natural Establishment of Specialist Plant Species after Topsoil Removal and Soil Perturbation in Degraded Calcareous Sandy Grassland

Specialist plant species in calcareous sandy grasslands are threatened by acidification and high nutrient levels in the topsoil. We investigated whether topsoil removal and soil perturbation in degraded sandy grasslands could lead to establishment of specialist species belonging to the threatened xeric sand calcareous grassland habitat. Restoration actions performed in 2006 resulted in increased soil pH and reduced nitrogen availability. We found early colonisztion of the perennial key species Koeleria glauca after both deep perturbation and topsoil removal, and high seedling establishment in topsoil removal plots 5 and 6 years following the restoration treatment (2011–2012). After topsoil removal, overall vegetation composition in 2012 had developed toward the undegraded community, with target species accounting for 20% of the community after topsoil removal, compared to 30% in the undegraded vegetation, and less than 1% in untreated controls. Deep perturbation led to 7% target species, while there were almost no effects of shallow perturbation 6 years following treatment. These results demonstrate that topsoil removal can promote colonization of target species of calcareous sandy grassland and highlights the importance of considering the regeneration niche for target species when implementing restoration measures .     

Nutrient limitation in boreal rich‐fen vegetation: A fertilization experiment

Abstract. Rich‐fen vegetation influenced by hay‐making in the Sølendet Nature Reserve, Central Norway, was fertilized with N, P and K in a full‐factorial fertilization experiment to investigate the nutrient limitation of plant growth at both community and species levels. Above‐ground biomass, shoot density and nutrient concentration were measured in several species and groups of species at three sites after two years of fertilization. At the community level, the results indicate multiple limitation by N and P in the two least productive rich‐fen communities: one characterized by small sedges and herbs, and the other by high abundance of Menyanthes trifoliata and tall sedges. Increased nutrient availability had no effect on a more highly productive, tall‐growing, spring‐influenced community, indicating no nutrient limitation. The results at the species level correspond well with those at the community level, indicating multiple limitation by N and P in most of the dominant and sub‐dominant species. However, P seems to limit growth more than N in Succisa pratensis, and N seems to limit growth more than P in Carex panicea. Furthermore, Eriophorum angustifolium seems to be limited by K. The results did not show which nutrient limits the growth of Carex dioica, C. lasiocarpa and Trichophorum cespitosum. Indications that growth in low‐productive, boreal rich‐fen communities is generally limited by P was not confirmed.     

Low Atmospheric Nitrogen Loads Lead to Grass Encroachment in Coastal Dunes,but Only on Acid Soils

The impact of atmospheric N-deposition on succession from open sand to dry, lichen-rich, short grassland, and tall grass vegetation dominated by Carex arenaria was surveyed in 19 coastal dune sites along the Baltic Sea. Coastal dunes with acid or slightly calcareous sand reacted differently to atmospheric wet deposition of 5–8 kg N ha⁻¹ y⁻¹. Accelerated acidification, as well as increased growth of Carex and accumulation of organic matter, was observed only at acid sites with pH_NaCl of the parent material below 6.0. At sites with slightly calcareous parent material, increased N-deposition had no effect. A trigger for grass encroachment seems to be high acidification in early successional stages to below pH_NaCl 4.0. Metals like Al or Fe become freely available and may hamper intolerant species. At acid sites, N-mineralization increases with elevated N-deposition, which may further stimulate Carex arenaria. Due to high growth plasticity, efficient resource allocation and tolerance of high metal concentrations, C. arenaria is a superior competitor under these conditions and can start to dominate the dune system. Carex-dominated vegetation is species-poor. Even at the moderate N-loads in this study, foliose lichens, forbs and grasses were reduced in short grass vegetation at acid sites. Species indicating these first effects of atmospheric deposition on dry, lichen-rich, short grasslands are identified and recommendations for restoration of grass-encroached sites given.     

Classification of habitats highlights priorities for conservation policies: The case of Spanish Mediterranean tall humid herb grasslands

The Mediterranean grasslands of Molinio-Holoschoenion are one of the lesser known natural habitat types of interest for conservation in the European Community. A study on their content with regard to their plant-communities and interest for plant conservation is conducted on a regional scale for the first time. A new comprehensive classification is proposed for this habitat type in Spain (Iberian Peninsula and Balearic and Canary Islands) where they have their highest European diversity. Twenty-five plant-community types are floristically identified by k-means clustering. Cluster analysis reveals a major hierarchical aggregation in two vegetation groups: (i) rush meadows of Scirpoides holoschoenus (L.) Soják; and (ii) communities of herbs and forbs dominated by Molinia caerulea (L.) Moench subsp. arundinacea (Schrank) H. Paul and/or Schoenus nigricans L. These groups are associated with different positions along a soil moisture gradient where rush formations occupy the positions farthest from the water table. Cluster Analysis also shows an uneven distribution of species with conservation interest. The highest content in endangered species is found in halophilous rushes. Grasslands on mineralised soils host the highest number of vulnerable species. Spanish Molinio-Holoschoenion plant communities were ranked using three criteria (regional responsibility, local rarity and habitat vulnerability) in order to set conservation priorities. The highest conservation values were achieved by coastal rush communities developing on dune slacks, and by forbs and rushes in the Baetic System growing on banks of streams with oligotrophic waters.     

Effects of a nutrient pulse supply on nutrient status of the Mediterranean trees Quercus ilex subsp. ballota and Pinus halepensis on different soils and under different competitive pressure

Nutrient availability is a key factor in Mediterranean ecosystems that affects the primary productivity and the community structure. The great variability of its natural availability is now increasing due to frequent fires, pollution events and changes in rainfall regime associated to climate change. Quercus ilex ssp. ballota and Pinus halepensis are the most abundant tree species in the NW Mediterranean basin. They frequently compete in the early and middle successional stages. We investigated the effects of N and P pulse supplies on nutrient uptake capacity in these two species in an after-fire field area and in nursery conditions on different soil types and competing conditions. In the field, N fertilisation had weak effects on nutrient concentration and mineralomass likely as a consequence of this nutrient not being limiting in this field site whereas P fertilisation increased the P mineralomass and the Mg, S, Fe, K and Ca concentrations and mineralomass in the different biomass fractions of both species 1 and 3 years after fertilisation application. In the nursery experiments, P fertilisation increased the mineralomass and concentrations of P, Mg, S, Fe, K and Ca in all biomass fractions including the roots in both species and in different soils and competition conditions. The increment of nutrient mineralomass was due to both the increase of growth and of nutrient concentrations. Both species were able to absorb significant amounts of the P applied by fertilisation (between 5 and 20%) in short time (18 months). Competing vegetation decreased the positive effects of P fertilisation, and in many cases the negative effect of competing vegetation on nutrient mineralomass was stronger when P availability was increased by fertilisation. Q. ilex subsp ballota showed a greater competitive ability for P than the more pioneer species Pinus halepensis in the field but not in the nursery conditions. Pinus halepensis had greater nutrient mineralomass in calcareous than in siliceous soils. Q. ilex subsp. ballota had a higher root biomass allocation and root nutrient allocation than P. halepensis, but both species showed a high capacity to increase their nutrient uptake when its availability increased by fertilisation, thus assuring a great nutrient reserve for future growth periods and contributing to retain nutrients in the ecosystem.     

Vertical canopy structure of Dutch chalk grasslands in relation to their management

Analysis of leaf canopy overtopping relationships was carried out using a non-destructive point quadrat method followed by a destructive stratified harvest of the above-ground phytomass in Dutch chalk grasslands with different management: summer sheep grazing and annual autumn mowing. The two methods of analysis are compared: e.g. relative leaf overtopping can be recorded by the point quadrat method but it is obscured in vertical vegetation profiles based on stratified phytomass distribution. However the stratified harvest method describes the relationship between canopy phytomass and light microclimate, recorded by measuring Photosynthetically Active Radiation (PAR) at different heights in the vegetation. Plant growth form during peak standing crop is of greater significance than Raunkiaerian life form in determining structure of chalk grassland vegetation. In annually mown grasslands, the tall graminoid growth form, shown by Brachypodium pinnatum, reduces PAR in the lower canopy and suppresses most other growth forms except those which can reach the higher canopy levels, e.g. clumped herbs such as Origanum vulgare. In contrast, grazing can result in a reduction of dominance from the tall graminoids and reduction of the abundance of taller, grazing-sensitive herbs, e.g. clumped and climbing herbs such as Origanum and Lathyrus pratensis, and an increase in grazing-tolerant species, e.g. smaller rosette herb growth forms, e.g. Leontodon hispidus and shorter rhizomatous or stoloniferous graminoids, e.g. Carex flacca and Briza media. Changes in both the overtopping hierarchy at the peak of the growing season and the intensity of overtopping in the course of a growing season are of conclusive importance in determining the relative abundance of species in the vegetation.     

甘肃中东部地区9种薹草属植物分布区群落特征和土壤养分状况

为探究薹草属(Carex L.)植物在不同植被类型中的分布状况,该研究对甘肃中东部地区9种薹草属(Carex L.)植物分布区的群落特征进行调查,并对土壤养分状况进行比较分析,以揭示野生薹草群落物种多样性和分布特征与土壤环境因子间的关系。结果表明：(1) 9种薹草群落物种多样性差异性较大,Shannon-Wiener多样性指数(H)、Simpson优势度指数(D_si)均以青绿薹草群落最高,亚柄薹草最低;Patrick丰富度指数(R)以异穗薹草群落最高,细叶薹草群落最低;Pielou均匀度指数(J_sw)以凹脉薹草群落最高,亚柄薹草最低。(2) 9种野生薹草属植物适宜生长的土壤pH呈中性或弱碱性,且有机质、氮素、钾素含量较丰富,磷含量偏低;土壤有机质、pH、全氮、全磷、全钾、碱解氮、速效磷、速效钾含量的平均值分别为41.07 g·kg^-1、8.35、1.16 g·kg^-1、0.65 g·kg^-1、5.60 g·kg^-1、47.94 mg·kg^-1、5.82 mg·kg^-1和100.60 mg·kg^-1。(3) 9种薹草属植物群落物种多样性与全氮、全磷、全钾、碱解氮、有机质、降雨量和海拔呈正相关关系,而与土壤pH、速效磷、速效钾呈负相关关系,且降雨量、土壤pH、速效磷和有机质对9种野生薹草属植物群落物种多样性影响较大。     

Nutrient-limited productivity of calcareous versus fleshy macroalgae in a eutrophic,carbonate-rich tropical marine environment

The results of a study of nutrient enrichment with nitrogen (N) and phosphorus (P) on productivity and calcification of fleshy and calcareous algae are reported in this study. Plants were collected from a nearshore eutrophic site in the Florida Keys (USA) and experimentally pulsed during the night with combinations of N and P. After several days of pulsing (7–10 days), net productivity, calcification, and alkaline phosphatase activity (APA), were measured. Productivity of fleshy algae were frequently enhanced by N, P, and N+P, during both summer and winter. Phosphorus limited the productivity of Hydroclathrus clathratus during winter and Ulva spp. during summer, whereas nitrogen limited the productivity of Laurencia intricata during both seasons. During summer, Dictyota cervicornis productivity was not enhanced by any nutrient enrichment. Nitrogen limited the productivity of the three calcareous species Penicillus capitatus, Penicillus dumetosus and Halimeda opuntia during winter and that of H. opuntia during summer. Neither N nor P enrichment increased calcification of calcareous species, and P enrichment greatly inhibited calcification of P. dumetosus during winter. Nutrient enrichment enhanced the productivity of the fleshy species to a greater extent than that of calcareous algae. The seawater DIN:SRP molar ratio was low at our eutrophic study site (molar ratio average of 3:1 during winter and 9:1 during summer) compared to more oligotrophic sites in the Florida Keys, suggesting that in carbonate-rich environments, eutrophication shifts nutrient regulation of productivity from P to N. APA activities of fleshy macroalage were higher than calcareous algae, and rates of all macro algae were 2- to 7-fold higher in summer compared to winter. Productivity was also about 3-fold higher in fleshy compared to calcareous species and about 2-fold higher in summer compared to winter. These results suggest that nutrient enrichment enhances productivity of fleshy algae to a greater extent than that of calcareous algae. Thus, overgrowth of calcareous algae by more opportunistic fleshy forms could reduce carbonate accretion in tropical coastlines experiencing increased eutrophication.