Invasive plants and their escape from root herbivory: a worldwide comparison of the root-feeding nematode communities of the dune grass <Emphasis Type="Italic">Ammophila arenaria</Emphasis> in natural and introduced ranges

Invasive plants generally have fewer aboveground pathogens and viruses in their introduced range than in their natural range, and they also have fewer pathogens than do similar plant species native to the introduced range. However, although plant abundance is strongly controlled by root herbivores and soil pathogens, there is very little knowledge on how invasive plants escape from belowground enemies. We therefore investigated if the general pattern for aboveground pathogens also applies to root-feeding nematodes and used the natural foredune grass Ammophila arenariaas a model. In the late 1800s, the European A. arenariawas introduced into southeast Australia (Tasmania), New Zealand, South Africa, and the west coast of the USA to be used for sand stabilization. In most of these regions, it has become a threat to native vegetation, because its excessive capacity to stabilize wind-blown sand has changed the geomorphology of coastal dunes. In stable dunes of most introduced regions, A. arenaria is more abundant and persists longer than in stabilized dunes of the natural range. We collected soil and root samples and used additional literature data to quantify the taxon richness of root-feeding nematodes on A.␣arenaria in its natural range and collected samples from the four major regions where it has been introduced. In most introduced regions A. arenaria did not have fewer root-feeding nematode taxa than the average number in its natural range, and native plant species did not have more nematode taxa than the introduced species. However, in the introduced range native plants had more feeding-specialist nematode taxa than A. arenaria and major feeding specialists (the sedentary endoparasitic cyst and root knot nematodes) were not found on A. arenaria in the southern hemisphere. We conclude that invasiveness of A. arenaria correlates with escape from feeding specialist nematodes, so that the pattern of escape from root-feeding nematodes is more alike escape from aboveground insect herbivores than escape from aboveground pathogens and viruses. In the natural range of A. arenaria, the number of specialist-feeding nematode taxa declines towards the margins. Growth experiments are needed to determine the relationship between nematode taxon diversity, abundance, and invasiveness of A. arenaria.     

Some considerations of and perspectives in coastal ecology

Ecophysiological studies of plant species in wet saline habitats have been one-sided in the past, emphasizing sodium chloride only. More emphasis should be given to anaerobiosis, to mechanical factors, to the effects of a surplus of other nutrients in seawater and coastal ecosystems and their metabolic impact as well as to aspects of nutrient (re)allocation. Research should incorporate theoretical models on plant strategies maximizing yield, minimizing risk. At the population level, a new operational parameter for ecological optima and ecological stress is presented considering normal and skewed distributions of plant parameters in relation to growth and the concentration of mineral nutrients.Some considerations on the ecological effects of phenotypic plasticity and genetic variability are given by emphasizing the aspects of diaspores.Future investigations of coastal vegetation should consider the interaction of populations within plant communities, including the regulating effects of animals.Nomenclature of vascular plants follows H. Heukels & R. van der Meijden, 1983. Flora van Nederland, 20th ed. Wolters-Noordhoff, Groningen; that of plant communities V. Westhoff & A. J. den Held, 1969. Plantengemeenschappen in Nederland. Thieme & Cie, Zutphen.     

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.     

Deer do not affect short‐term rates of vegetation recovery in overwash fans on Fire Island after Hurricane Sandy

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Does salt stress constrain spatial distribution of dune building grasses Ammophila arenaria and Elytrichia juncea on the beach?

Rising sea levels threaten coastal safety by increasing the risk of flooding. Coastal dunes provide a natural form of coastal protection. Understanding drivers that constrain early development of dunes is necessary to assess whether dune development may keep pace with sea‐level rise. In this study, we explored to what extent salt stress experienced by dune building plant species constrains their spatial distribution at the Dutch sandy coast. We conducted a field transplantation experiment and a glasshouse experiment with two dune building grasses Ammophila arenaria and Elytrigia juncea. In the field, we measured salinity and monitored growth of transplanted grasses in four vegetation zones: (I) nonvegetated beach, (II) E. juncea occurring, (III) both species co‐occurring, and (IV) A. arenaria dominant. In the glasshouse, we subjected the two species to six soil salinity treatments, with and without salt spray. We monitored biomass, photosynthesis, leaf sodium, and nutrient concentrations over a growing season. The vegetation zones were weakly associated with summer soil salinity; zone I and II were significantly more saline than zones III and IV. Ammophila arenaria performed equally (zone II) or better (zones III, IV) than E. juncea, suggesting soil salinity did not limit species performance. Both species showed severe winter mortality. In the glasshouse, A. arenaria biomass decreased linearly with soil salinity, presumably as a result of osmotic stress. Elytrigia juncea showed a nonlinear response to soil salinity with an optimum at 0.75% soil salinity. Our findings suggest that soil salinity stress either takes place in winter, or that development of vegetated dunes is less sensitive to soil salinity than hitherto expected.     

A double-edged sword: parental care increases risk of offspring infection by a maternally vectored parasite

Parental care can protect offspring from predators but can also create opportunities for parents to vector parasites to their offspring. We hypothesized that the risk of infection by maternally vectored parasites would increase with the frequency of mother–offspring contact. Ammophila spp. wasps (Hymenoptera: Sphecidae) build nests in which they rear a single offspring. Ammophila species exhibit varied offspring provisioning behaviours: some species enter the nest once to provision a single, large caterpillar, whereas others enter the nest repeatedly to provision with many smaller caterpillars. We hypothesized that each nest visit increases the risk of offspring parasitism by Paraxenos lugubris (Strepsiptera: Xenidae), whose infectious stages ride on the mother wasp (phoresy) to reach the vulnerable Ammophila offspring. We quantified parasitism risk by external examination of museum-curated Ammophila specimens—the anterior portion of P. lugubris protrudes between the adult host''s abdominal sclerites and reflects infection during the larval stage. As predicted, Ammophila species that receive larger numbers of provisions incur greater risks of parasitism, with nest provisioning behaviour explaining ca 90% of the interspecific variation in mean parasitism. These findings demonstrate that parental care can augment, rather than reduce, the risk of parasite transmission to offspring.     

Etudein situ de l'influence de l'humidité et de la teneur en nitrate d'un sol dunaire sur l'accumulation et la réduction du nitrate chez l'oyat (Ammophila arenaria L.)

Résumé L'étude du fonctionnement du cycle de l'azote dans les milieux dunaires des c?tes méditerranéennes fran?aises a conduit à analyser le comportement de l'oyat,Ammophila arenaria, au cours de son développement en réponse aux variations de l'humidité et de la teneur en nitrate du sol. Les mesures d'activité nitrate réductase et de la teneur en nitrate dans les feuilles ont été effectuées simultanément avec les mesures des teneurs en eau et en nitrate du sol. Cette plante a des activités nitrate réductase ne dépassant pas 0,27 μmoles h⁻¹ g⁻¹ matière fra?che, qui n'ont pu être mesuréesin vivo qu'en présence de nitrate exogène dans le milieu d'incubation. Il appara?t que la plante réduit ou accumule préférentiellement le nitrate selon que l'on se situe en période humide ou sèche. Pour des valeurs voisines ou inférieures à 2% d'humidité dans les vingt premiers centimètres de sol, l'activité nitrate réductase décro?t et des quantités importantes de nitrate s'accumulent (169 μg N.NO₃.g⁻¹ matière sèche). Dans ces conditions, le nitrate dans la solution du sol peut atteindre une concentration élevée (6 meq.l⁻¹). Inversement, au-delà de 2% d'humidité le nitrate endogène s'épuise et l'activité nitrate réductase augmente. La régulation de l'assimilation et de l'accumulation du nitrate par l'oyat dépend de son alimentation hydrique et done des quantités de nitrate absorbées à partir de la solution du sol et véhiculées dans la sève brute.

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In situ study of the influence of dune soil humidity and nitrate contents on the nitrate accumulation and reduction in marram (Ammophila arenaria L.)

Summary The search for a better understanding of the nitrogen cycle in the sand dunes of french mediterranean beaches have driven us to study the habits ofAmmophila arenaria. In this way, we have followed its responses to different soil water potentials and nitrate contents of the soil during its development. Nitrate reductase activities and nitrate contents of leaves have been measured simultaneously with water and nitrate contents of soil. This plant has a slight nitrate reduction activity, not more than 0,27 μmoles h⁻¹ g⁻¹ fresh matter, which could be measuredin vivo only by the addition of nitrate to the incubation media. It seems that the plant reduces or else it stores, in a preferential way the nitrate whether it is in a humid or dry period. If humidity values are near or below 2%, the marram diminishes its assimilation activity and stores important amounts of nitrate (169 μg N−NO₃ ⁻.g⁻¹ dry matter). In these conditions the nitrate can be highly concentrated in the soil solutions (6 meq.l⁻¹). Opposite, above 2% of humidity, the nitrate reductase activity decreases and the endogenous nitrate is consumed. The regulation of the assimilation and storage of nitrate by the marram is mainly a function of the soil humidity and so, of the amount of nitrate taken from the soil solution and carried into the xylem.

     

Colonization of the root zone ofAmmophila arenaria by harmful soil organisms

The role of harmful soil organisms in the degeneration ofAmmophila arenaria at coastal foredunes was examined by the growing of seedlings ofA. arenaria in soil samples collected from its root zone. Three sites, each representing a successive stage in foredune succession were examined: (1) a highly mobile dune (sand accretion of 80 cm year⁻¹) with vigorousA. arenaria, colonizing only the upper 30-cm of the annually deposited layer of sand, (2) a mobile dune with vigorousA. arenaria (sand accretion of 22 cm year⁻¹) and a 1-metre soil profile completely colonized by roots and (3) a stable dune (no sand accretion) with degeneratedA. arenaria and young roots mainly present in the upper 0–10 cm. In the upper part of the highly mobile site, the presence of harmful soil organisms was confined to the root layers and at the mobile site for all depth layers a significant growth reduction ofA. arenaria was observed due to the activity of harmful soil organisms. At the stable site, however, growth had only been reduced in some of the depth layers. At all sites newly formed roots ofA. arenaria had been colonized by harmful soil organisms within one year. If present in sand prior to root growth harmful soil organisms reduced root length and root hair formation severely and they enhanced branching of the roots. It is concluded that harmful soil organisms initiate degeneration ofA. arenaria in stable dunes by attack of the root system, which makes the plants suffer from abiotic stress.     

Mycorrhizae and succession in plantings of beachgrass in sand dunes

A survey of arbuscular mycorrhizal fungi (AMF), arbuscular mycorrhizae (AM), and hyphal networks of AMF was carried out in sand dune sites of different successional stages in the Province Lands Area of Cape Cod National Seashore, Massachusetts. The study focused on large-scale plantings (each of 12–20 ha) of American beachgrass (Ammophila breviligulata) aged 0–7 yr and five adjacent natural dune areas. Sample sites ranged in vegetative cover from barren to forested. Spores of 17 species of AMF were recovered from the dunes. Over the successional sequence, there were increases in the richness and spore populations of the AMF community, the extent of colonization of A. breviligulata roots, and the mycorrhizal inoculum potential of the soil. Unvegetated sites lacked propagules of AMF, but roots of planted culms of A. breviligulata (which carried propagules of AMF) became mycorrhizal in <1 yr after planting. Spores were recovered from previously AMF-free sites that had been planted with beachgrass for 47 wk, and five species of AMF sporulated in sites <6 yr old. Significant hyphal networks were not present in any of the planted areas (<6 yr old at the time of sampling), but did occur in natural areas. The rate of invasion of areas planted to A. breviligulata by later successional plant species may in part depend upon the establishment of a vigorous network of hyphae of AMF in a site.