Soil pH and species diversity in coastal dunes

Soil pH was measured at two different spatial scales in coastal dunes on Norderney, North Sea, and in Mecklenburg-Vorpommern, Baltic Sea, Germany. Relationships between the variability in soil pH, species richness and species diversity are presented. Species richness and diversity were highest in grey dunes, where soil pH was at intermediate levels; both variables were lower in yellow and brown dunes. The variability in pH increased with increasing species diversity and also with scale. Overall, soil pH variability decreased with increasing vegetation cover. The lowest pH heterogeneity was found in heath dominated by Empetrum nigrum L. and grey dunes dominated by Campylopus introflexus (Hedw.) Brid. Increasing abundance of dominant species and decreasing species diversity of vegetation apparently reduces soil heterogeneity. Decreasing species diversity of vegetation is likely to explain decreasing variability in soil pH.     

Impact of anthropogenic disturbances on beetle communities of French Mediterranean coastal dunes

In coastal dunes, influenced by anthropogenic activities such as tourism, it is important to determine the relative influence of environmental factors at different spatial scales to evaluate the sensitivity of local communities to disturbances. We analyzed beetle communities of 14 dunes of the French Mediterranean coast: four in the relatively preserved Camargue area, and ten in the Var department, where tourism is intensive. Beetle communities were studied three times in early spring using sand sampling. Species-environment relationships were evaluated at the regional, landscape and local scale using redundancy analysis (RDA) and variability partitioning. About 28 species were identified, of which 15 were sand-specialist species, which accounted for more than 93% of total abundance. The beetle communities of Camargue were significantly different from those of the Var department owing to the pullulation of a Tenebrionid species (Trachyscelis aphodioides Latr.) in the Var, except for one restored dune where the community was very similar to those of Camargue. Our results showed no longitudinal gradient between the two regions. Local factors (dune height, preservation and disturbance index) significantly explained most of the variation in the dominance of T. aphodioides, while some other local factors were important for other psammophilous species. This study also suggests that dune beetle communities are strongly affected on beaches intensively managed for tourism, but beetles are still abundant in much disturbed sites.     

Fluctuating deposition of ocean water drives plant function on coastal sand dunes

Sea‐level rise will alter the hydrology of terrestrial coastal ecosystems. As such, it becomes increasingly important to decipher the present role of ocean water in coastal ecosystems in order to assess the coming effects of sea‐level rise scenarios. Sand dunes occur at the interface of land and sea. Traditionally, they are conceived as freshwater environments with rain and ground water as the only water sources available to vegetation. This study investigates the possibility of ocean water influx to dune soils and its effect on the physiology of sand dune vegetation. Stable isotopes are used to trace the path of ocean water from the soil to the vegetation. Soil salinity, water content and δ¹⁸O values are measured concurrently with stem water and leaf tissue of eight species during the wet and dry season and from areas proximal and distal to the ocean. Our results indicate the dune ecosystem is a mixed freshwater and marine water system characterized by oceanic influence on dune hydrology that is spatially heterogeneous and fluctuates temporally. Ocean water influx to soil occurs via salt spray in areas 5–12 m from the ocean during dry season. Accordingly, vegetation nearest to the sea demonstrate a plastic response to ocean water deposition including elevated integrated water use efficiency (δ¹³C_leaf) and uptake of ocean water that comprised up to 52% of xylem water. We suggest physiological plasticity in response to periodic ocean water influx may be a functional characteristic common to species on the leading edge of diverse coastal habitats and an important feature that should be included in modeling coastal ecosystems. Rising sea level would likely cause a repercussive landward shift of dune species in response to encroaching maritime influences. However, human development would restrict this process, potentially causing the demise of dune systems and the protection from land erosion they provide.     

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

Question: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump‐backed relationship between species number and shrub cover? Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney. Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m² size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen). Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump‐backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump‐backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co‐variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa. Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.     

Soil nutrients trump intraspecific effects on understory plant communities

Understanding the links between intraspecific genetic variation and patterns of diversity in associated communities has been the primary focus of community genetics or ‘genes-to-ecosystem’ research in ecology. While other ecological factors, such as the abiotic environment, have well-documented influences on communities, the relative contributions of genetic variation versus the environment to species interactions remains poorly explored. In this study, we use a common garden experiment to study a coastal dune plant community dominated by the shrub, Baccharis pilularis, which displays a morphological dimorphism in plant architecture. We found the differences in the understory plant community between erect and prostrate morphs of Baccharis to be statistically significant, but small relative to the impacts of nutrient additions (NPK and C additions), for the richness, cover, and biomass of the understory plant community. There were no significant interactions between Baccharis morphology and nutrient-addition treatments, suggesting the influence of nutrient addition was consistent between erect and prostrate morphs. Moreover, we found no difference in overall plant community composition between Baccharis morphs, while NPK additions led to shifts in understory community composition compared to unfertilized shrubs. In sum, our results indicate that nutrients are the more important factor governing understory plant community structure in a coastal dunes ecosystem followed by intraspecific variation in dominant shrub architecture. Our results address a growing call to understand the extended consequences of intraspecific variation across heterogeneous environments in terrestrial ecosystems.     

Darwin's naturalization hypothesis: scale matters in coastal plant communities

Darwin proposed two seemingly contradictory hypotheses for a better understanding of biological invasions. Strong relatedness of invaders to native communities as an indication of niche overlap could promote naturalization because of appropriate niche adaptation, but could also hamper naturalization because of negative interactions with native species (‘Darwin's naturalization hypothesis’). Although these hypotheses provide clear and opposing predictions for expected patterns of species relatedness in invaded communities, so far no study has been able to clearly disentangle the underlying mechanisms. We hypothesize that conflicting past results are mainly due to the neglected role of spatial resolution of the community sampling. In this study, we corroborate both of Darwin's expectations by using phylogenetic relatedness as a measure of niche overlap and by testing the effects of sampling resolution in highly invaded coastal plant communities. At spatial resolutions fine enough to detect signatures of biotic interactions, we find that most invaders are less related to their nearest relative in invaded plant communities than expected by chance (phylogenetic overdispersion). Yet at coarser spatial resolutions, native assemblages become more invasible for closely‐related species as a consequence of habitat filtering (phylogenetic clustering). Recognition of the importance of the spatial resolution at which communities are studied allows apparently contrasting theoretical and empirical results to be reconciled. Our study opens new perspectives on how to better detect, differentiate and understand the impact of negative biotic interactions and habitat filtering on the ability of invaders to establish in native communities.     

Phenological and biogeographical aspects of coastal dune plant communities in southern Brazil

The coastline of southern Brazil is characterized by vast sand dunes. From March 1982 to April 1984 the dune vegetation was sampled along a transect between the Atlantic shore and an inland freshwater marsh, and phenological cycles of plant species were observed at monthly intervals.The occurrence of 23 annual species in the local dunes and peak germination, growth and flowering of 21 perennial species during spring, summer and fall, suggest that unfavourable temperatures and inundation of slacks during winter are seasonally limiting factors for this flora.Differences in the floristic composition between this region and its southern and northern extremes in the southwestern Atlantic together with a marked seasonality of the total species number and of germination and flowering periods, suggest that this area corresponds to a warm temperate biogeographic transition zone between northern tropical and southern cold temperate regions.     

Species-richness and soil phosphorus in plant communities in coastal New South Wales

Species-richness (recorded as number of species in 16 m² quadrats), in more than 60 coastal sites developed over various substrata, exhibited a significant non-linear relationship with soil total phosphorus extracted from associated soil samples. Although the general form of this relationship is not incompatible with a resource competition model, the significant proportion of the total variance accounted for by the species-richness/phosphorus relationship suggests that a more detailed examination of phosphorus use at a community level will be profitable.     

Ectomycorrhizal fungal communities on seedlings and conspecific trees of Pinus mugo grown on the coastal dunes of the Curonian Spit in Lithuania

Ectomycorrhizal (ECM) communities of mature trees and regenerating seedlings of a non-native tree species Pinus mugo grown in a harsh environment of the coastal region of the Curonian Spit National Park in Lithuania were assessed. We established three study sites (S1, S2, and S3) that were separated from each other by 15 km. The ECM species richness was rather low in particular for mature, 100-year-old trees: 12 ectomycorrhizal taxa were identified by molecular analysis from 11 distinguished morphotypes. All 12 taxa were present on seedlings and on mature trees, with between 8–11 and 9–11 taxa present on seedlings and mature trees, respectively. Cenococcum geophilum dominated all ECM communities, but the relative abundance of C. geophilum mycorrhizas was nearly two times higher on seedlings than on mature trees. Mycorrhizal associations formed by Wilcoxina sp., Lactarius rufus, and Russula paludosa were also abundant. Several fungal taxa were only occasionally detected, including Cortinarius sp., Cortinarius obtusus, Cortinarius croceus, and Meliniomyces sp. Shannon’s diversity indices for the ECM assemblages of P. mugo ranged from 0.98 to 1.09 for seedling and from 1.05 to 1.31 for mature trees. According to analysis of similarity, the mycorrhizal communities were similar between the sites (R = 0.085; P = 0.06) and only slightly separated between seedlings and mature trees (R = 0.24; P < 0.0001). An incidental fruiting body survey that was conducted weakly reflected the below-ground assessment of the ECM fungal community and once again showed that ECM and fruiting body studies commonly supply different partial accounts of the true ECM fungal diversity. Our results show that P. mugo has moved into quite distinct habitats and is able to adapt a suite of ECM symbionts that sufficiently support growth and development of this tree and allow for natural seedling regeneration.     

Resource heterogeneity generated by shrubs and topography on coastal sand dunes

In early stages of primary succession, colonizing plants can create resource patches that influence the abundance and distribution of other species. To test whether different colonizing shrubs generate contrasting patches on coastal sand dunes, we compared soil characteristics and light availability under the nitrogen-fixing shrub Lupinus arboreus, under the non-nitrogen-fixing shrub Artemisia pycnocephala, and between shrubs on dunes at a site in northern California. Concentrations of inorganic nitrogen and net nitrogen mineralization rates were generally 1–10 times greater in soil under Lupinus than under Artemisia or between shrubs. Soil water content was mostly lower under shrubs. Mean photon flux density near ground level was reduced by at least 80% at 35 cm inside shrub canopies. Topography appeared to have more effect on soil moisture but less direct effect on nitrogen availability than did Lupinus. However, Lupinus probably increases nitrogen levels more on higher, drier dunes. Microhabitats under and between nitrogen-fixing shrubs constitute a mosaic of individually poor but complementary patches in which high levels of light and moderate levels of soil nitrogen are present but tend not to occur together.     

Dune slack restoration in Dutch mainland coastal dunes

The 45,000 ha Dutch dunes are relatively well protected but they suffer from severe desiccation. During the last decade a lot of knowledge has been obtained about the steering processes of eco-hydrology in dunes resulting in well-founded schemes to restore wet dune slack communities. In two case studies a large-scale restoration of the landscape was necessary. Since 1987 in the North-Holland Dune Reserve the groundwater catchment has been reduced (from 8 million m³ to 2 million m³ per year) and the resulting development monitored. In another case, large-scale intervention on 35 ha of the Amsterdam Water Supply Dunes in 1995 resulted in the restoration of active sand drift with opportunities for the establishment of pioneer vegetation. In small parts of the area, the surface has stabilised as a result of vegetation growth, but elsewhere drifting sand has spread and covered previously stable surfaces. Many wet to moist dune slacks developed and results relating to geomorphology, hydrology and vegetation are presented. Finally, in 1996 yet another project of 50 ha was carried out in the Meijendel dunes, in which infiltration ponds were removed to allow vegetation succession to develop from scratch. Results of these three case studies will be presented. If only hydrological restoration is carried out, sod-cutting was found to be necessary to restore the whole series of vegetation types of dune slacks. In two cases an area was denuded. The seed bank and surrounding slacks seem to play an important role in the success of restoration, this is especially true for the pioneer vegetation types. Future monitoring will prove whether critical (Red List) plant species will return.     

Water relations of coastal plant communities near the ocean/freshwater boundary

Summary Salinity and isotope ratios were determined in water from several wells in the Florida Keys, and tidal inlets. Both D/H and ¹⁸O/¹⁶O ratios of water from wells and tidal inlets were highly correlated to their salinity. Water from standing pools was enriched in deuterium and oxygen-18 relative to their salinity because of evaporation processes. ¹⁸O/¹⁶O and D/H ratios of stem water from plants of several different communities at Sugar Loaf Key, ranging from hardwood hammocks to mangroves, were highly correlated to their predawn water potential. The correlation was consistent with the presence of high salinity in waters with high ¹⁸O and D content. Most individuals from each community were either utilizing water with isotopic characteristics typical of freshwater or of ocean water, while only a few individuals had stem water with isotopic ratios intermediate to these two water sources.     

Soil microorganisms control plant ectoparasitic nematodes in natural coastal foredunes

Belowground herbivores can exert important controls on the composition of natural plant communities. Until now, relatively few studies have investigated which factors may control the abundance of belowground herbivores. In Dutch coastal foredunes, the root-feeding nematode Tylenchorhynchus ventralis is capable of reducing the performance of the dominant grass Ammophila arenaria (Marram grass). However, field surveys show that populations of this nematode usually are controlled to nondamaging densities, but the control mechanism is unknown. In the present study, we first established that T. ventralis populations are top-down controlled by soil biota. Then, selective removal of soil fauna suggested that soil microorganisms play an important role in controlling T. ventralis. This result was confirmed by an experiment where selective inoculation of microarthropods, nematodes and microbes together with T. ventralis into sterilized dune soil resulted in nematode control when microbes were present. Adding nematodes had some effect, whereas microarthropods did not have a significant effect on T. ventralis. Our results have important implications for the appreciation of herbivore controls in natural soils. Soil food web models assume that herbivorous nematodes are controlled by predaceous invertebrates, whereas many biological control studies focus on managing nematode abundance by soil microorganisms. We propose that soil microorganisms play a more important role than do carnivorous soil invertebrates in the top-down control of herbivorous ectoparasitic nematodes in natural ecosystems. This is opposite to many studies on factors controlling root-feeding insects, which are supposed to be controlled by carnivorous invertebrates, parasitoids, or entomopathogenic nematodes. Our conclusion is that the ectoparasitic nematode T. ventralis is potentially able to limit productivity of the dune grass A. arenaria but that soil organisms, mostly microorganisms, usually prevent the development of growth-reducing population densities.     

Germination responses to abiotic stress shape species distributions on coastal dunes

Plant species occupy distinct zones on coastal dunes, but the mechanisms limiting their distributions have not been fully explained. We combined field surveys of plant distributions and abiotic conditions with controlled germination experiments to assess the contribution of germination requirements to plant zonation. Species presence and abiotic conditions were measured in ten transects across the barrier dune at Waquoit Bay, Massachusetts. Germinating seeds of six species were exposed to four fully crossed treatments: pre-treatment (soaked in fresh water, soaked in salt water, or not soaked), temperature (low, moderate, or high), soil salinity (none, moderate, or high), and light (full light, shade, dark). Species distributions in the field were affected by both distance from the shore and presence of dominant shrubs. Germination tolerance of soil salinity reflected species zonation: species found on the front slope of the dune tolerated salinity, while germination of other species was limited by salinity alone or by salinity in combination with high temperature. Shrubs reduced soil surface temperature and decreased light, but these conditions had limited effects on germination. These results indicate that limitations to germination can contribute to explaining species distributions on coastal dunes.     

Effects of flooding, salinity and herbivory on coastal plant communities, Louisiana, United States

Flooding and salinity stress are predicted to increase in coastal Louisiana as relative sea level rise (RSLR) continues in the Gulf of Mexico region. Although wetland plant species are adapted to these stressors, questions persist as to how marshes may respond to changed abiotic variables caused by RSLR, and how herbivory by native and non-native mammals may affect this response. The effects of altered flooding and salinity on coastal marsh communities were examined in two field experiments that simultaneously manipulated herbivore pressure. Marsh sods subjected to increased or decreased flooding (by lowering or raising sods, respectively), and increased or decreased salinity (by reciprocally transplanting sods between a brackish and fresh marsh), were monitored inside and outside mammalian herbivore exclosures for three growing seasons. Increased flooding stress reduced species numbers and biomass; alleviating flooding stress did not significantly alter species numbers while community biomass increased. Increased salinity reduced species numbers and biomass, more so if herbivores were present. Decreasing salinity had an unexpected effect: herbivores selectively consumed plants transplanted from the higher-salinity site. In plots protected from herbivory, decreased salinity had little effect on species numbers or biomass, but community composition changed. Overall, herbivore pressure further reduced species richness and biomass under conditions of increased flooding and increased salinity, supporting other findings that coastal marsh species can tolerate increasingly stressful conditions unless another factor, e.g., herbivory, is also present. Also, species dropped out of more stressful treatments much faster than they were added when stresses were alleviated, likely due to restrictions on dispersal. The rate at which plant communities will shift as a result of changed abiotic variables will determine if marshes remain viable when subjected to RSLR. Received: 8 April 1998 / Accepted: 15 June 1998