Facilitation of seedling establishment by an endemic shrub in tropical coastal sand dunes

Facilitation is predicted to occur on coastal sand dunes as these ecosystems have the harsh physical conditions common during primary succession. In a tropical dune system on the Gulf of Mexico the spatial patterns of plant distribution were analyzed and the hypothesis that facilitation is the responsible mechanism was tested by monitoring changes in the micro-environment and performing a seedling planting experiment under natural conditions. Densities of seedlings and adults of late colonizer grasses were two to six times greater beneath the early colonizer shrub, Chamaecrista chamaecristoides, than in the exposed areas. Temperatures on the sand surface, wind speed and sand accretion were significantly reduced by the shade of Chamaecrista. Only phosphate contents during the dry season were significantly higher in sand in the shade than in exposed sand. In the field experiment, successful establishment of seedlings of two late colonizer grasses (Trachypogon plumosus and Schizachyrium scoparium) was low but was concentrated exclusively beneath the shrubs. The introduced plants were reproductive one year after onset of the experiment and remained vigorous after three years. The observed spatial aggregation among the target species may be a result of improved conditions in the shade of the shrub, which facilitated the survival and establishment of late colonizers. On top of the environmental amelioration, protection from recurrent disturbances such as substrate mobility is an additional beneficial effect of the shrub.     

A test for hydrotropic behavior by roots of two coastal dune shrubs

Root hydrotropism could be a means by which plants forage for limited and patchy distributions of soil water. While root hydrotropism has been induced in distinctly artificial conditions, it is unclear if it operates in natural settings. Here, we tested for this possibility in seedlings of two species of dune shrubs. Growth of individual roots in sand-filled observation chambers was monitored in response to moisture-rich patches and resultant soil water gradients. Chambers were designed so that roots could intercept the moisture gradients but not the moisture-rich patches simply through gravitropism. While up to 12% of the Eriogonum parvifolium roots grew into the moisture-rich patches, comparable root growth was observed in the control. None of the Artemisia californica roots grew into the patches. Thus, in a reasonable simulation of field conditions, we found no compelling evidence for hydrotropic root behavior in seedlings of these two dune shrubs. Our results leave the ecological significance of root hydrotropism in question.     

Adaptations enhancing survival and establishment of seedlings on coastal dune systems

Studies on the survival and establishment of seedlings in coastal and lacustrine sand dune systems suggest that nutrient deficiency, lack of moisture, sand accretion, salt spray and predation are probably the most important limiting factors. Seedlings employ both avoidance and tolerance strategies to with-stand the different stresses. For example, seedling recruitment of dune species coincided with periods of high moisture availability and occurred in years with high well distributed rainfall. The seedlings exhibited rapid vertical elongation of roots, avoided moisture stress by leaf rolling, dense hair on leaves, fleshy cotyledonary leaves, and by growing in the shade of nurse plants. A certain proportion of seedlings of all species survived partial burial and showed stimulation of leaf growth, total leaf area, number of tillers and total dry matter. Seedlings of most species could not survive complete burial, however, some species did grow through the sand deposit by an increase in the number of nodes and length of internodes. Plants responded to salt spray by inhibition of seed germination, developing resistance, increase in leaf thickness and reduced uptake of Na and Cl ions. Several characters such as higher vigour of seedlings, larger seed size and seed polymorphism were also of adaptive significance to the species.     

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.     

The influence of food supplementation on a coastal dune rodent community

Studies of the rodent community in regenerating coastal dune forest areas north of Richards Bay, South Africa, have indicated that a number of species occupy newly rehabilitating areas. Unstable environmental conditions result in habitat changes, rather than competitive interactions, being the major determinant of rodent community development. This led to the present study, which investigated the effects of supplementary food on rodent assemblages. We attempted to stabilize food availability in order to elucidate some of the mechanisms responsible for the development of the rodent community. By only increasing an existing portion of the resource spectrum, we reduced species diversity, probably through dominance by competitive superior exploiters of the augmented resource. Thus, species diversity decreased with food supplementation, as a result of an increase in the abundance of Mastomys natalensis rather than because of a change in the number of species or the abundance of other species.     

Post-fire regeneration strategies of Californian coastal sage shrubs

Summary Regeneration methods for coastal sage srub vegetation after fire were studied in the coastal Santa Monica Mountains of southern California. Six sites were sampled two years after a large fire of fall, 1978. The intensity of fire varied. Foliar cover and flowering incidence were recorded for individuals regenerating by resprouting or from seed. Resprouting plants contributed most to post-fire recovery, comprising 95% of the relative foliar shrub cover; 84% of resprout and 47% of seedling cover had flowered. An ANOVA of reproductive mode and fire intensity indicates that resprout total cover and individual size are significantly greater than those of seedlings, regardless of fire intensity. Among sites the average foliar cover of resprouts exceeded that of seedlings by factors ranging from 9 to 63. All coastal sage species examined resprout, although the potential vigor of resprouting appears to vary widely within genera (e.g. Encelia, Eriogonum, and Salvia) and even within species. In the second growing season following fire seedling density increased due to seeds shed by resprouted shrubs. Most of the cover on these stands of coastal sage scrub is destined to be either crown-sprouted individuals or their progeny.     

Restoration of coastal dune slacks in the Netherlands

In order to stop the continuous decline of typical dune slack communities along the Dutch coast, restoration projects have been carried out since 1952. Restoration measures consisted of re-introducing traditional management techniques in dune slacks, such as mowing, grazing and sod removal, or constructing artificial dune slacks to compensate for lost biodiversity elsewhere. An analysis of successful and unsuccessful projects showed that constructing new dune slacks was not very successful for maintaining new populations of endangered dune slack species, since such projects were often carried out in areas where seed banks were depleted, while hydrological conditions and seed dispersal mechanisms were sub-optimal. The construction of sand dikes to prevent sea intrusion in large beach plains was, unintentionally, a temporary success for the establishment of many Red List species, although such measures often disrupted natural dune slack formation. Successful sites were all characterised by a regular discharge of calcareous groundwater provided by local or regional hydrological systems, where not very long ago populations of typical dune slack plants were present. Under such conditions, sod removal was a successful measure to create pioneer stages which were relatively stable, due to a very slow accumulation of organic matter in the topsoil. It is argued that new and more flexible coastal defence strategies can provide new opportunities for natural and relatively stable pioneer stages of dune slack formation, suitable for the long term preservation of endangered dune slack species.     

Plant-scale pattern among herbs and shrubs of a fire-dominated coastal plain forest

Herb and shrub layer vegetation was surveyed in the oak-pine upland forest of the Pine Barrens region of New Jersey (USA). To describe spatial distributions of individual species, 15 4 m² quadrats were censused in each of 34 stands. At the scale of whole stands, stem densities were significantly related to several forms of historical disturbance including canopy defoliation and fire. At the scales of 1 m² and 4 m² quadrats, distributions of the 19 most abundant species were all aggregated to some degree, with clonal species more strongly clustered than non-clonal. Shrub stems tended to be strongly clustered, and to be distributed independently of leaf litter. By contrast, tree seedlings were only marginally clustered, and were significantly more common in quadrats with a lower litter cover. Clonal herbs exceeded shrubs in degree of aggregation, but were also sensitive to the local accumulation of leaf litter. Distributions of non-clonal herbs were sensitive to leaf litter, but were only marginally clustered, and appeared to depend upon gaps in the tree canopy. Rather than supporting a single life history, a combination of disturbance types has allowed a variety of species with different ecological requirements and growth forms to coexist in the same site. Overall species diversity can be maintained in this ecosystem without invoking community-level interactions.     

Responses to invasion and invader removal differ between native and exotic plant groups in a coastal dune

The spread of exotic, invasive species is a global phenomenon that is recognized as a major source of environmental change. Although many studies have addressed the effects of exotic plants on the communities they invade, few have quantified the effects of invader removal on plant communities, or considered the degree to which different plant groups vary in response to invasion and invader removal. We evaluated the effects of an exotic succulent, iceplant (Carpobrotus edulis), on a coastal dune plant community in northern California, as well as the community responses to its removal. To assess possible mechanisms by which iceplant affects other plants, we also evaluated its above- and belowground influences on the germination and growth of a dominant exotic annual grass, Bromus diandrus. We found that iceplant invasion was associated with reduced native plant cover as well as increased cover and density of some exotic plants—especially exotic annual grasses. However, iceplant removal did not necessarily lead to a reversal of these effects: removal increased the cover and density of both native and exotic species. We also found that B. diandrus grown in iceplant patches, or in soil where iceplant had been removed, had poorer germination and growth than B. diandrus grown in soil not influenced by iceplant. This suggests that the influence of iceplant on this dune plant community occurs, at least in part, due to belowground effects, and that these effects remain after iceplant has been removed. Our study demonstrates the importance of considering how exotic invasive plants affect not only native species, but also co-occurring exotic taxa. It also shows that combining observational studies with removal experiments can lead to important insights into the influence of invaders and the mechanisms of their effects.     

Alien plants of coastal dune habitats in southern Spain

Alien plants from psammophilous ecosystems have been studied in southern Spain. Sixteen sites, which are included in the Andalusian Network of Protected Spaces, have been chosen along 1100 km of Andalusian coasts in order to verify the degree of abundance and influence of these species on different communities. Out of the 26 localized species, the increase of some invasive plants, such as Arctotheca calendula, Agave americana, A. sisalana, Cortaderia selloana, Lantana camara and Opuntia dillenii, affects the conservation of some natural coastal ecosystems.     

Status of coastal dune communities invaded by Chrysanthemoides monilifera

It has often been assumed that the introduced species Chrysanthemoides monilifera is displacing native species in invaded coastal communities in Australia but no direct evidence has been presented that this displacement is occurring and, if so, to what extent. At an invaded site near Moruya, New South Wales, we found strong negative correlations between the presence (measured as cover, frequency or volume of shrub) of C. monilifera and Acacia longifolia, the hitherto dominant native species, and of C. monilifera and Lomandra longifolia. The negative correlations are partly accounted for by differences in the amounts of each species on different zones on the dune, but the negative correlations remain even within these zones. Also a comparison with uninvaded sites indicates that C. monilifera is displacing A. longifolia from the fore-dune and mid-dune but is having less effect in the swale. In addition, there is a trend towards reduced seed production of A. longifolia when growing in proximity to C. monilifera. There is no significant difference in the presence of L. longifolia in the uninvaded and invaded sites. In invaded sites, significantly fewer C. monilifera seedlings occur in the presence of L. longifolia. Thus it appears that the observed negative associations arise from different causes. Chrysanthemoides monilifera is displacing the structurally similar, and previously dominant A. longifolia, while the tussock-forming L. longifolia is locally hindering the invasion.     

Factors controlling soil development in sand dunes: evidence from a coastal dune soil chronosequence

Aerial photographs, maps and optically stimulated luminescence dates were combined with existing soil data to construct high resolution chronosequences of soil development over 140 years at a temperate Atlantic UK dune system. Since soil formation had progressed for varying duration under different climate and nitrogen deposition regimes, it was possible to infer their relative influence on soil development compared with location-specific variables such as soil pH, slope and distance to the sea. Results suggest that soil development followed a sigmoid curve. Soil development was faster in wet than in dry dune habitats. In dry dunes, rates were greater than in the literature: they increased with increasing temperature and nitrogen deposition and decreased with increasing summer gales. The combination explained 62% of the variation. Co-correlation meant that effects of nitrogen deposition could not be differentiated from temperature. In wet dune habitats rates increased with temperature and decreased with gales. The combination explained only 23% of the variation; surprisingly, rainfall was not significant. Effects of location-specific variables were not significant in either habitat type. Nitrogen accumulation was faster in wet than dry dune habitats, averaging 43 kg N ha^?1 per year overall. Nitrogen accumulation greatly exceeded inputs from atmospheric deposition, suggesting rates of input for biological N fixation are 10–60 kg N ha^?1 per year. Recent climate and/or nitrogen deposition regimes may have accelerated soil development compared with past rates. These data suggest the importance of changing climate on soil development rates and highlight the contribution of biological N fixation in early successional systems.     

Environmental tuning of an insect ensemble: The tenebrionid beetles inhabiting a Mediterranean coastal dune zonation

Few studies have investigated insect ensembles, i.e. phylogenetically bounded groups of species that use a similar set of resources within a community. The zonation of dune vegetation makes these ecosystems ideal for the study of insect ensembles in a short space. In this study, we investigated if the tenebrionid beetles forming an ensemble on a dune zonation showed variations in community organization (relative abundances and species diversity) in different but spatially associated biotopes defined by different plant communities. Three biotopes (corresponding to European Commission habitat 2110, 2120 and 2230) of a well-preserved Mediterranean dune were sampled using square plots of 2 × 2 m at three places. To investigate if there was some association between species and habitat we applied a χ² test. Variations in community structure parameters were investigated using Shannon index. The three biotopes host tenebrionid communities with similar species composition and overall abundances, confirming that they form a single ensemble. However, tenebrionid species are differently associated with different biotopes along the zonation, with some species occurring with different proportions among the biotopes. A local selection process can be postulated as a mechanism responsible for these differences.     

Climate and coastal dune vegetation: disturbance, recovery, and succession

The sand dune habitats found on barrier islands and other coastal areas support a dynamic plant community while protecting areas further inland from waves and wind. Foredune, interdune, and backdune habitats common to most coastal dunes have very different vegetation, likely because of the interplay among plant succession, exposure, disturbance, and resource availability. However, surprisingly few long-term data are available describing dune vegetation patterns. A nine-year census of 294 plots on St. George Island, Florida suggests that the major climatic drivers of vegetation patterns vary with habitat. Community structure is correlated with the elevation, soil moisture, and percent soil ash of each 1 m² plot. Major storms reduce species richness in all three habitats. Principle coordinate analysis suggests that changes in the plant communities through time are caused by climatic events: changes in foredune vegetation are correlated with temperature and summer precipitation, interdune vegetation with storm surge, and backdune vegetation with precipitation and storm surge. We suggest that the plant communities in foredune, interdune, and backdune habitats tend to undergo succession toward particular compositions of species, with climatic disturbances pushing the communities away from these more deterministic trajectories.     

The importance of facilitation in the zonation of shrubs along a coastal salinity gradient

Question: What are the interactive roles of abiotic stress and plant interactions in mediating the zonation of the shrub Tamarix chinensis along a salinity gradient? Location: Yellow River estuary (37°46′N, 119°09′E), northeast China. Methods: We surveyed the zonation of T. chinensis along a salinity gradient and quantified its salt tolerance using a pot experiment. In two field experiments, we transplanted T. chinensis seedlings into salt marsh, transitional zone and upland habitats, manipulated neighbours and quantified survivorship and biomass to examine neighbour effects. We also quantified vegetation effects on abiotic conditions in each zone. Results: Tamarix chinensis dominated the transitional zone, but was absent in upland and salt marsh habitats. In the pot experiment, T. chinensis grew well in freshwater treatments, but was inhibited by increasing salinity. Field experiments revealed that competition from neighbours limited T. chinensis growth in the uplands, while T. chinensis transplants were limited, with or without neighbours, in the salt marsh by high soil salinity. In the transitional zone, however, T. chinensis transplants performed better with than without neighbours. Vegetation removal significantly elevated soil salinity in the transitional zone, but not in other zones. Conclusions: Competition, facilitation and abiotic stress are all important in mediating the zonation of T. chinensis. Within its physiological stress tolerance range, or fundamental niche, it is limited by plant competition in low salinity habitats, and facilitated by neighbours in high salt stress habitats, but cannot survive in salt marshes having salinities above its salt stress tolerance limit. Our results have implications for understanding the relationships between facilitation and stress gradients.     

Stomatal responses to humidity of coastal and interior populations of a Californian shrub

Summary Plants of two populations of Diplacus aurantiacus, a subshrub of the Californian chaparral, were compared for their stomatal response to water vapor concentration gradients. Plants of a coastal and an interior population were compared when grown under both low and high humidities. When grown at high humidity the coastal plants exhibited higher conductances and higher transpiration/photosynthesis ratios at all leaf-to-air water vapor concentration gradients than did the interior plants. Although all of the plants examined showed a pronounced stomatal response to humidity the response did not result in the degree of regulation of water-use efficiency reported for other Californian coastal species.     

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.     

Intraspecific competition between shrubs in a semi-arid savanna

Tree-on-tree competitive interactions may be more important in affecting the distribution of the tree components of savannas than inter-specific competition with grasses. The presence of intraspecific competition is expected to negatively affect inter-tree spacing, individual size distributions and plant physiology as well as survival/mortality. In this field removal experiment on Acacia mellifera, one of South Africa’s most common encroachers on nutrient-poor soils, the growth, water relations and mortalities of shrubs where all neighbouring woody competitors were removed (target) were monitored three times during each of three growing seasons. After 3 years, the nitrogen and carbon isotopic ratios of the study plants were analysed. Target shrubs benefitted from removal of neighbours, resulting in greater growth, less water stress, a relatively small degree of canopy dieback and reduced reliance on N₂ fixation. Target shrubs grew by 25 ± 4% in height relative to 7 ± 4% for controls, with the targets suffering a maximum of <15% canopy dieback compared to up to 60% in the controls. Severe environmental stress is known to affect neighbour interactions among shrubs and competition may constrain shrub sizes and avoid density-dependent mortality. In contrast, release from competition in our study may have allowed greater growth of target plants, increasing their total evapo-transpirational leaf surface areas and leaving them vulnerable to drought and water stress. Intratree competition on shallow nutrient-poor soils in savannas may thus aid the persistence of bush encroachment by regulating the sizes of individual shrubs below the threshold of drought vulnerability.