Sandblasting as a possible factor controlling the distribution of plants on a coastal dune system

Intensity of the abrasive effect of wind-borne sand – sandblasting – in addition to other environmental factors was measured at two vegetation zones on a sandy beach and one site at an inland area. One zone on the beach included foredunes sparsely vegetated by dune species such as Carex kobomugi and Calystegia soldanella. The other zone which was located ∼50 m inland from the first zone was flat grassland dominated by inland species such as Miscanthus sinensis and Imperata cylindrica var. Koenigii. The inland site consisted of short grassland located 3 km inland from the beach. Intensity of sandblasting was estimated by the whiteness of a transparent plastic sheet exposed to the air for 2 weeks. This sheet turned whitely opaque when it was abraded by wind-borne sand. The other environmental factors measured at the beach were intensity of salt spray, soil water content, soil salinity, and sand accumulation, while intensity of salt spray was the only additional factor measured at the inland site. Intensity of sandblasting was considerably higher at the foredune zone, while that at the grassland zone was as low as that at the inland site. Considerable salt spray was detected at the foredune and grassland zones. Differences in other environmental factors were small between the two zones on the beach. In order to compare the difference in tolerance to sandblasting, a jet of sand was applied to one ordinary species, C. kobomugi, from the foredune and two species, M. sinensis and I. cylindrica, from the grassland zone. The difference in tolerance was determined by the decrease in the area of green leaf after applying sandblasting with commercial sandblaster and/or spraying with sea water. M. sinensis and I. cylindrica lost much of the leaf area after sandblasting and salt spraying, while C. kobomugi lost little. These results indicated that one of the characteristic environmental factors of a foredune is the high intensity of sandblasting accompanied by salt spray, and that species found in the foredune are more tolerant to sandblasting than species distributing in more inland areas.     

Zonation of vegetation on lacustrine coastal dunes: effects of burial by sand

More than 50 years ago it was proposed that zonation of major plant species on coastal dunes was determined by salt spray. Here, we argue against this hypothesis because (i) salt concentrations rarely exceed toxic levels; (ii) high precipitation in temperate latitudes washes the salt off the plants; (iii) major salt spray events occur in late autumn and winter when plants are dormant; and (iv) zonation also occurs on lacustrine dunes. Instead, we show evidence that zonation may be caused by burial because plant distribution was correlated with sand deposition and species were eliminated when burial exceeded their limits of tolerance, thus creating zones of different plant species. We conclude that in temperate regions (i) burial by sand may be among the most important factors in zonation, while salt spray may play a secondary role; and (ii) single environmental factors cannot be the determinants of a phenomenon as complex as species zonation.     

Shoreline processes and establishment of Phragmites australis in a coastal plain estuary

Phragmites australis occurs extensively along undisturbed salt-marsh shorelines of Delaware Bay. The species has been considered indicative of human disturbance when found in estuarine marshes in the USA. It is suggested that geomorphic processes associated with coastal submergence provide an analog of human disturbances which can enable Phragmites australis to become established naturally. Deposition of sand bodies (or rafted debris) can suppress existing vegetation and allow Phragmites to become established. Subsequently, even if the sand or debris is moved, erosional truncation of the intertidal profile can inhibit recolonization by the original dominant shoreline species, Spartina alterniflora.     

Ecophysiological adaptations of coastal halophytes from foredunes and salt marshes

Ecophysiological strategies of coastal halophytes from foredunes and salt marshes are discussed. A comparison is made of the factors that limit growth in salt marshes and sand dunes. In salt marshes, zonation and succession are primarily governed by variation in soil salinity, which strongly depends on inundation with seawater. Results are described of experiments which aim at separating salinity and inundation effects on growth, osmotic and mineral relations in a comparison of salt-marsh halophytes. The growth response of plants cannot simply be correlated (and causally explained) with the concentration of Na, Cl, and K in the tissues. Also, the compatible osmotic solutes proline and methylated quaternary ammonium compounds may accumulate both in species with a positive response to increased salinity and in species with a growth reduction under seawater inundation. More likely inadequate adaptation of the plants water potential with these components is partly the cause of retarded growth. Disfunctioning of the plant in this respect may be at three levels: (a) total water potential of the plant, (b) (loss) of turgor pressure potential; (c) regulation at the cellular level. The ecological importance of some factors in seawater other than sodium chloride is considered. In coastal sand dunes airborne rather than soil salinity limits plant growth, together with the effects of abrasion, sand accretion, drought and the poor nutrient status of the dune sand. Adaptations of sand-dune species to these factors may consist of: large seeds with storage tissue germinating in the dark and seedling growth enough to emerge through the accreted sand. Aerial parts must be resistant to mechanical damage (high wind speed and abrasion), possibly by a sclerophyllous and tough structure. Efficient nutrient uptake, translocation and retranslocation seem to help survive sand-dune species in a nutrient-poor rooting medium.     

Responses of photosynthesis,leaf conductance and growth to different salinities in three coastal dune plants

The effects of soil-water salinity on growth and photosynthesis of three coastal dune plants were examined by salt-treatment in order to clarify the causal relationship between salinity and plant distribution in a dune habitat. Plants were cultivated hydroponically at three salinity levels: 0, 10 and 100 mM NaCl. With the 100 mM salt treatment,Calystegia soldanella (C₃ species) had the highest relative growth rate (RGR) (0.085 g g⁻¹ d⁻¹), followed byCarex kobomugi (C₃) (0.066), andIschaemum anthephoroides (C₄) (0.060). This order coincides with the distribution pattern of the three species on coastal dunes;Calystegia soldanella is generally distributed in more seaward areas whereasI. anthephoroides occurs further inland. The order of RGR was determined exclusively by leaf area ratio (LAR) among the three species. Due to its C₄ pathway,I. anthephoroides had higher net photosynthetic rate (Pn) and net assimilation rate (NAR) than the two C₃ plants at all NaCl concentrations, despite its low RGR. This apparent discrepancy is explainable by differences of LAR among the three species; LAR ofI. anthephoroides was lowest, and about half that ofCalystegia soldanella. These results suggest that LAR is one of the main determinants of salt tolerance based on RGR, whereas Pn or NAR may not be significant. This article is dedicated to Professor Hideo Iwaki, University of Tsukuba, in appreciation of the sincere encouragement he has given to the authors.     

Biological soil crusts and their microenvironment: Impact on emergence,survival and establishment of seedlings

To elucidate the impact of biological soil crusts (BSCs) on the establishment of habitat-typical vascular plant species, we studied the effects of seed location (surface versus sub-surface), age of crusts (initial versus stable), long rainy periods (continuous versus discontinuous watering) and microenvironment (cracks versus no cracks). In addition, we investigated growth height, phytomass and N-content of one vascular plant species (Phleum arenarium). Initial crusts were compared with older, stable crusts using seven habitat-typical plant species representing different life forms (annuals versus perennials). Our model ecosystem, situated in the temperate zone (but edaphically dry), is characterised by calcareous sand with threatened pioneer vegetation (Koelerion glaucae). We carefully translocated soil monoliths of these crusts and analysed the effects under common garden conditions.     

Limiting factors for seedling emergence and establishment of the invasive non-native Rosa rugosa in a coastal dune system

The relative importance of seed, habitat and microhabitat limitation has rarely been investigated for invasive non-native species, although this is critical for their effective management and for predicting future range expansion. Rosa rugosa is an abundant non-native shrub in coastal habitats of NW Europe; it is common along the Baltic coast but more scarce in exposed dunes of the North Sea. To investigate whether invasion of exposed dunes is limited by seed, habitat and microhabitat limitation, seedling emergence and establishment were examined in a factorial sowing, transplant and disturbance experiment. Twenty plots were randomly placed in each of five dune habitats (white dune, Empetrum dune, grey dune, outer dune heath, inner dune heath), and studied over 2 years. Seedling emergence in control subplots was zero in all habitats, whereas 96% and 98% of the undisturbed and disturbed seeded subplots produced seedlings. Disturbance had a positive effect on emergence and subsequent survival in white dune, outer and inner dune heath. Seedling survival and growth, and growth of transplanted seedlings, were markedly lower in grey dune. These findings indicate that establishment of R. rugosa is seed-limited in coastal dune habitats, and that the species is able to establish in both active and fixed dunes once seeds have arrived. Although differential seedling emergence and growth indicate that habitats differ in their degree of invasion susceptibility by R. rugosa, the positive influence of small-scale disturbance suggests microhabitat limitation in some dune habitats as well. Dune management should aim to reduce seed production and dispersal of R. rugosa in near-natural sites, and anthropogenic changes of habitat dynamics should be prevented.     

Germination and reproductive biology in salty conditions of Asplenium marinum (Aspleniaceae), a European coastal fern

Although there are several species of ferns that are important components of many coastal ecosystems, the adaptations that allow these species to live in salty conditions have been studied only in two Acrostichum species. We have studied the effects of salt on germination, development of the gametophyte and sexual expression of the fern Asplenium marinum, which lives on coastal cliffs of Europe. Cultures at four different levels of salt concentration, 0%, 0.98%, 1.96% and 3.26%, were established for this purpose. Salt caused a significant delay in spore germination, and prevented it completely at high concentrations (3.26%). However, spores were able to recover germination ability after a period of seawater incubation once salinity concentrations decreased. Variable salinity conditions are a typical feature of sea cliffs where halophytes can grow. A salty culture medium also affects the development of the prothalli. Higher salinity of the substrate produced a significant decrease in gametophyte sizes, and affected the development of gametangia as well. Only males were found in a 0.98%-salt medium, and no gametangia were observed in cultures at 1.96%-salt medium. Recovered spores after one and a half month in high salinity concentration, 3.26%, were able, when transplanted to 0%-salt medium, to develop normal sexual prothalli, and this occurred in a shorter time than spores without any prior treatment. A. marinum can be considered as a halophyte fern that is able to germinate and grow in salty conditions. But it takes advantage of rainy periods that cause decrease of salt concentration, which seems to be important in particular for gametangia development and fertilization.     

Abiotic constraints on the competitive ability of exotic and native grasses in a Pacific Northwest prairie

In prairie ecosystems, abiotic constraints on competition can structure plant communities; however, the extent to which competition between native and exotic plant species is constrained by environmental factors is still debated. The objective of our study was to use paired field and greenhouse experiments to evaluate the competitive dynamics between two native (Danthonia californica and Deschampsia cespitosa) and two exotic (Schedonorus arundinaceus and Lolium multiflorum) grass species under varying nutrient and moisture conditions in an upland prairie in the Willamette Valley, Oregon. We hypothesized the two invasive, exotic grasses would be more competitive under high-nutrient, moderate-moisture conditions, resulting in the displacement of native grasses from these environments. In the field, the experimental reduction of competition resulted in shorter, wider plants, but only the annual grass, Lolium multiflorum, produced more aboveground biomass when competition was reduced. In the greenhouse, the two exotic grasses produced more total biomass than the two native grasses. Competitive hierarchies were influenced by nutrient and/or moisture treatments for the two exotic grasses, but not for the two native grasses. L. multiflorum dominated competitive interactions with all other grasses across treatments. In general, S. arundinaceus dominated when in competition with native grasses, and D. cespitosa produced the most biomass in monoculture or under interspecific competition with the other native grass, D. californica. D. californica, D. cespitosa, and S. arundinaceus all produced more biomass in high-moisture, high-nutrient environments, and D. cespitosa, L. multiflorum, and S. arundinaceus allocated more biomass belowground in the low nutrient treatment. Taken together, these experiments suggest the competitive superiority of the exotic grasses, especially L. multiflorum, but, contrary to our hypothesis, the native grasses were not preferentially excluded from nutrient-rich, moderately wet environments. Laurel Pfeifer-Meister and Esther M. Cole contributed equally to this work.     

Facilitation and sand burial affect plant survival during restoration of a tropical coastal sand dune degraded by tourist cars

Coastal sand dunes support various ecosystem services, including storm protection and tourism. Restoration programs are often critical to preserve this ecosystem due to its fragility and high degree of degradation. Dune restoration still suffers from a general lack of knowledge of the ecological processes controlling tropical dune communities. We investigated if facilitation can increase restoration success by assisting plant survival in a Brazilian coastal dune degraded by buggies at the Environmental Protection Area of Jenipabu, RN, Brazil. We performed two field experiments on dune crest sites degraded by buggies. The first experiment tested how the presence of established vegetation and coconut mesh, mimicking soil‐stabilizing effect of vegetation, facilitates seedling survival and establishment of the early successional dominant dune plant Canavalia maritima (Fabaceae). The second experiment tested if coconut mesh and initial irrigation would allow the establishment of C. maritima outside the vegetation, using both seeds and transplants. We found that the presence of established vegetation positively affected seedling survival. Application of mesh positively affected microclimatic conditions and slightly increased survival, but only for seeds. Initial irrigation only had a small effect on transplanted seedlings survival. Nevertheless, there was extreme seedling mortality in the experiment in particular from sand burial. We conclude that restoration of these dune crests is very difficult. Facilitation by vegetation can reduce plant mortality by sand burial and desiccation, but successful restoration is likely to require a major effort involving a very high number of seedlings or seeds.     

The distribution and ecology of coastal species on roadsides

The invasion of roadsides by coastal species is a response to the use of de-icing salt. The most widespread species on Britain's roadsides, Puccinellia distans, occurs in the north and east of England and Wales. This distribution is due to the varying use and effects of de-icing salt which are related in turn to regional differences in climate and traffic density. On roadsides, seed dispersal is greater in the direction of traffic flow and the smaller seeds of Puccinellia distans are carried further than the larger seeds of Plantago maritima. Species on roadsides occur in distinct zones with the coastal species confined to the saline road margin. The causes of this zonation were investigated by examining seedling establishment in experimental plots. Seedlings were only able to establish in grass plots when the salinity was sufficiently high to kill the competing glycophytic grasses. On bare soil, the halophytes established and grew satisfactorily when salt was added but grew very poorly in the absence of salt. After two years Plantago maritima overcame this apparent requirement for salt on bare soil but Puccinellia distans and Spergularia marina did not.Acknowledgements: N. E. Scott was in receipt of a N.E.R.C. Studentship. The authors wish to thank the members of the B.S.B.I. and the Biological Records Office for information on species distribution.     

The effects of water and multi-nutrient stress on xylem sap chemistry,photosynthesis and transpiration of seedlings of two Eucalypts

Summary Ten seedlings each of Eucalyptus kitsoniana Maiden and Eucalyptus globulus Labill. were subjected to two levels of water stress and two levels of nutrient stress (macro and micro-nutrients) in a greenhouse for 3 weeks. The objectives were to determine the degree to which seedlings show differences in sap chemistry, photosynthesis and transpiration that relate to the environments in which these two species live. Whole plants were then extracted for xylem sap using a pressure chamber and the sap was analyzed for 14 elements using an inductively coupled plasma spectrometer and a nitrometer. For E. kitsoniana water and nutrient stress, applied separately or in combination, significantly reduced leaf conductance, transpiration, photosynthesis and midday water potential. Nutrient stress alone had less effect than water stress on most functions measured. Water stress alone reduced the root/shoot ratio; the combination of water and nutrient stress increased the root/shoot ratio, primarily because of reduced shoot weight. In E. kitsoniana, water stress alone or in combination with nutrient stress increased the xylem sap concentrations of B and Si. Multi-nutrient stress alone, or in combination with water stress, significantly decreased sap Zn and K. For this species, sap N was decreased by nutrient stress, but increased by water stress. E. globulus had significantly lower transpiration rates and less root mass than E. kitsoniana. Slightly lower leaf conductance and photosynthesis were not significant in E. globulus compared to E. kitsoniana. Water and nutrient stress reduced conductance, transpiration (except for nutrient stress) and photosynthesis, and the effects of water stress on E. globulus were greater than the effects of nutrient stress. Midday water potential was reduced by water stress. Water or nutrient stress alone did not alter seedling root/shoot ratio, but the combination of water and nutrient stress significantly increased the root/shoot ratio for both species. For E. globulus, sap concentrations of Mn, Na, Si and K were increased by water stress (alone or in combination with nutrient stress). Sap N increased with water stress or combined stresses, but decreased under nutrient stress alone. When the two species were compared, E. globulus generally had lower or similar nutrient concentrations in the sap, with Ca, Mg, Mn and P significantly lower than in E. kitsoniana. Seedlings of these two species show strong site adaptations to water and nutrient availability.     

Physical and chemical effects of litter on plant establishment in semi-natural grasslands

A field experiment was conducted in a semi-natural grassland to study the interspecific variation in the effect of litter on seedling emergence and establishment and separate physical from chemical effects. Seeds of seven forb species were sown in plots subjected to either litter amendment (0, 400 or 900 g m⁻²) or water extracts of litter (corresponding to 400 and 900 g litter m⁻²). In addition, an extract was treated with activated carbon to estimate the possible effects of secondary chemical compounds. The response to plant litter differed amongst species: negative, neutral and positive responses were observed. Anthriscus sylvestris was the only species with a strong positive response to litter. We found no consistent relation between seed size and response to plant litter. Physical effects of litter were generally stronger than chemical effects. However, water extract of litter inhibited emergence in three species. Activated carbon removed the negative effect of the litter extract, which suggests that the effect was caused by an inhibitory chemical compound rather than by increased competition in response to nutrients added via the extract. The balance between facilitative and inhibitory effects of litter depended on species identity and litter quantity. Facilitative effects dominated at low and intermediate quantities of litter, and inhibitory effects at high litter quantities. One species, Campanula rotundifolia, showed a switch from positive to negative responses with increasing quantities of litter. However, we found no general threshold for litter quantity valid across species.     

Interactive effects of global and regional change on a coastal ecosystem

Vertical distribution and diel vertical migration of a zooplankton community were studied at two stations off Central Peru in April 2006. Zooplankton was collected at five depth strata by vertical hauls with Hydo-Bios multinet (300-μm mesh, 0.25-m² mouth size). The zooplankton community was distributed in relation to a strong, shallow oxycline (1 ml l^?1 oxygen isopleth generally above 36 m). The highest total abundance was always in the upper, well-oxygenated layer. The most important species were: Acartia tonsa (72.86%), Centropages brachiatus (7.5%), and Paracalanus parvus (3.1%); Acartia tonsa was the dominant species at all times. Larvae of the polychaete Magelona sp. (7.5%) and larvae of the brachiopod Discinisca lamellosa (3.5%) were numerically dominant in April and small copepods e.g. Oncaea venusta (3.88%) were numerically dominant during August. Five distinct patterns of vertical distribution and migration in relation to the oxygen minimum layer were distinguished in this study: (1) Ontogenetic vertical migration through the oxycline (Acartia tonsa adults, nauplii, and copepodids), (2) permanent limitation to layers above the oxycline (e.g. Oikopleura sp., most invertebrate larvae), (3) distribution mostly below the oxycline with occasional migration into the layers just above the oxycline (Eucalanus inermis), (4) Diel Vertical Migration (Centropages brachiatus), and (5) reverse Diel Vertical Migration (larvae of the polychaete Magelona sp.).     

Impact of grazing and atmospheric nitrogen deposition on the vegetation of dry coastal dune grasslands

Abstract. A five-year experimental study was carried out to examine the combined effects of grazing and atmospheric nitrogen deposition on the vegetation of three dry dune grasslands: one short species-rich, one short species-poor, and one predominated by tall graminoids. Additional fertilization with nitrogen had no significant effect, neither in grazed nor in non-grazed plots. Exclusion of grazing by rabbits resulted in an increase in the frequency of perennial graminoids and a decrease in the frequency of annual graminoids and herbs. Nevertheless, species diversity remained the same in the species-rich grassland. During the experiment, the above-ground biomass increased in all nongrazed plots and the amount of bare soil and mosses decreased. The vegetation changes occurred mainly within one year after the exclusion of grazing. An exception is the grass-dominated site where the amount of Calamagrostis epigejos increased gradually from ca. 20 % in the first two years to about 50 % in the fourth and fifth year. Grazing by rabbits seems essential to prevent graminoids to become predominant in the dry dunes. If graminoids are dominant, grazing by horses can be an appropriate method to restore the original grassland vegetation. After six months of grazing by horses the grass-dominated site showed a decrease of the frequency of perennial graminoids, from 95 % to 80 %, and an increase of the frequency of perennial herbs, from 2.5 % to between 13 and 20 %.     

Effects of grazing on seedling establishment: the role of seed and safe-site availability

Abstract. The first objective of this paper was to assess the effects of grazing on seedling establishment of two species whose relative abundance at the adult stage is affected by grazing in a contrasting fashion. Second, we evaluated the relative importance of seed versus safe-site availability in explaining the effect of grazing on seedling establishment. We monitored seedling establishment on a grazed area, on two areas which had not been grazed for two and seven years, and on plots which had been experimentally defoliated. The species compared were Dan-thonia montevidensis, a native perennial grass which dominates both grazed and ungrazed communities, and Leontodón taraxacoides, an invading exotic rosette species from the Compositae family. Continuous grazing enhanced seedling establishment of both species through its effect on the availability of safe sites. Seed availability accounted for only one, but very important, grazing effect: the lack of response by L. taraxacoides to the defoliation in the seven-year old exclosure. Its seed supply was depleted by exclusion of grazing and, consequently, its short-term regeneration capacity after disturbance was lost.     

Effects of emergence time on survival and growth in an early old-field plant community

Summary The time at which plants emerge from the soil is shown to be correlated with both survival and growth in each of four years in a plant community emerging after yearly plowing. For all seven species investigated, earlier emerging individuals generally had both a higher biomass and probability of survival. There were differences among species in the effect of emergence time on biomass, with the growth of upright, annual species being more suppressed by later emergence than the growth of other species. No significant differences were found among species in the effects of emergence time on survival. It was expected that differences among species in the effect of emergence time on fitness might lead to a correlation between the patterns of emergence time and species characteristics such as growth form or lifespan. Mean emergence times did vary significantly among both the seven species and the four years of the study. However, there was no correlation between emergence time and species lifespan, growth form, abundance, or competitive ability. The lack of a correlation between the selection pressure on species and their emergence time could be due low heritability, insufficient time for selection and evolution, and/or selection on correlated characters.     

Effects of exogenous putrescine on gas-exchange characteristics and chlorophyll fluorescence of NaCl-stressed cucumber seedlings

The effects of 10 mM putrescine (Put) treated by spraying on leaves on growth, chlorophyll content, photosynthetic gas-exchange characteristics, and chlorophyll fluorescence were investigated by growing cucumber plants (Cucumis sativus L. cv. ChangChun mici) using hydroponics with or without 65 mM NaCl as a salt stress. Salt stress caused the reduction of growth such as leaf area, root volume, plant height, and fresh and dry weights. Furthermore, net photosynthesis rate (P _n), stomatal conductance (g _s), intercellular CO₂ concentration (C _i), and transpiration rate (T _r) were also reduced by NaCl, but water use efficiency (WUE; P _n/T _r) showed a tendency to be enhanced rather than reduced by NaCl. However, Put alleviated the reduction of P _n by NaCl, and showed a further reduction of C _i by NaCl. The reduction of g _s and T _r by NaCl was not alleviated at all. The enhancement of WUE by NaCl was shown to have no alleviation at day 1 after starting the treatment, but after that, the enhancement was gradually reduced till the control level. Maximum quantum efficiency of PSII (F _v/F _m) showed no effects by any conditions based on the combination of NaCl and Put, and in addition, kept constant values in plants grown in each nutrient solution during this experimental period. The efficiency of excitation energy capture by open photosystem II (PSII) (F _v′/F _m′), actual efficiency of PSII (Φ_PSII), and the coefficient on photochemical quenching (qP) of plants with NaCl were reduced with time, and the reduction was alleviated till the control level by treatment with Put. The F _v′/F _m′, Φ_PSII, and qP of plants without NaCl and/or with Put showed no variation during the experiment. Non-photochemical quenching of the singlet excited state of chlorophyll a (NPQ) showed quite different manner from the others as mentioned above, namely, continued to enhance during the experiment.