Susana Rodríguez-Echeverría

Physiological integration increases the survival and growth of the clonal invader Carpobrotus edulis.

Clonal growth seems to be a common trait for many of the most aggressive invasive plant species. However, little research has been conducted to determine the role of clonality in the successful invasion of new areas by exotic species. Carpobrotus edulis (L.) N.E. Br. is a mat-forming succulent plant, native to South Africa that is invasive in coastal dunes of Australia, New Zealand, USA and Southern Europe. Although Carpobrotus edulis is a clonal plant, there is no information on the role of clonality for the invasion by this species, therefore the objective of this study was to test whether or not physiological integration improves the performance of C. edulis invading coastal sand dunes. To do that, a 6-month field experiment was designed in which the stolon connections between the apical ramets and the C. edulis mats were severed to prevent physiological integration. This treatment was applied to ramets growing under high and low competition with the native species. Apical ramets with intact stolon connections were used as control. Integration improved the survivorship and growth of apical ramets, both in high and low competition. Connected ramets showed a more pronounced increase of clonal growth (estimated as stolon length) during the experimental period and a higher total biomass and number of ramets at the completion of the experiment. In terms of survivorship, the benefit of integration was greater under high competition. Physiological integration can therefore be considered an important factor in the invasiveness of C. edulis, both in open space and in direct competition with the native plants.

Impact of wildfire return interval on the ectomycorrhizal resistant propagules communities of a Mediterranean open forest

Ectomycorrhizal (ECM) fungi, in particular their spores and other resistant propagules, play an important role in secondary succession processes that facilitate regeneration after disturbance events. In this study, the effects of high and low wildfire frequencies (respectively short and long fire return intervals) on the resistant propagules communities (RPCs) of a Mediterranean open pine forest were compared. Soil samples were collected in four mountain sites with different fire return intervals and used to test ectomycorrhiza development in two hosts, Pinus pinaster and Quercus suber. RPCs were characterized by direct sequencing of fungal internal transcribed spacer (ITS) regions from individual ECM root tips. Eighteen ECM species were detected in the bioassay. The most frequently found fungi were Cenococcum geophilum, Inocybe jacobi, Thelephora terrestris, Tomentella ellisii on both hosts and Rhizopogon luteolus and R. roseolus on maritime pine. A short fire return interval reduced the species richness of the ECM community found on Q. suber, promoted species like R. roseolus and reduced the abundance of other species (e.g. R. luteolus). The abundance of I. jacobi was positively affected by long fire return interval, but decreased significantly with recurrent fires. These results indicate that changes in fire frequency can alter the structure, composition and diversity of ECM communities, which could compromise the resilience of the ecosystem in highly disturbed areas.

Differential effectiveness of novel and old legume–rhizobia mutualisms: implications for invasion by exotic legumes

The degree of specialization in the legume–rhizobium mutualism and the variation in the response to different potential symbionts are crucial factors for understanding the process of invasion by exotic legumes and the consequences for the native resident plants and bacteria. The enhanced novel mutualism hypothesis predicts that exotic invasive legumes would take advantage of native rhizobia present in the invaded soils. However, recent studies have shown that exotic legumes might become invasive by using exotic introduced microsymbionts, and that they could be a source of exotic bacteria for native legumes. To unravel the role of novel and old symbioses in the progress of invasion, nodulation and symbiotic effectiveness were analyzed for exotic invasive plants and native co-occurring legumes in a Mediterranean coastal dune ecosystem. Although most of the studied species nodulated with bacteria from distant origins these novel mutualisms were less effective in terms of nodulation, nitrogenase activity and plant growth than the interactions of plants and bacteria from the same origin. The relative effect of exotic bradyrhizobia was strongly positive for exotic invasive legumes and detrimental for native shrubs. We conclude that (1) the studied invasive legumes do not rely on novel mutualisms but rather need the co-introduction of compatible symbionts, and (2) since exotic rhizobia colonize native legumes in invaded areas, the lack of effectiveness of these novel symbiosis demonstrated here suggests that invasion can disrupt native belowground mutualisms and reduce native legumes fitness.

Nematode interactions in nature : Models for sustainable control of nematode pests of crop plants?

van der Putten, W. H., Cook, R., Costa, S., Davies, K. G., Fargette, M., Freitas, H., Hol, W. H. G., Kerry, B. R., Maher, N., Mateille, T., Moens, M., de la Pena, E., Piskiewicz, A., Raeymaekers, A., Rodriguez-Echeverria, S., van der Wurff, A. W. G. (2006). Nematode interactions in nature: models for sustainable control of nematode pests of crop plants? Advances in Agronomy, 89, 227-260.

Diversity of AMF associated with Ammophila arenaria ssp. arundinacea in Portuguese sand dunes

Dune vegetation is essential for the formation and preservation of sand dunes and the protection of the coast line. Coastal sand dunes are harsh environments where arbuscular mycorrhizal fungi (AMF) play an important role in promoting plant establishment and growth. We present a study of the diversity of AMF associated with A. arenaria ssp. arundinacea in two locations of the Portuguese coast under a Mediterranean climate. These two locations were selected to compare a well-preserved dune system from a protected area with a degraded dune system from a public beach. AMF diversity was assessed mainly by cloning and sequencing of a fragment of the ribosomal SSU using the primer NS31 and AM1. Most of the 89 AMF clones obtained from the rhizosphere and roots of A. arenaria belonged to the genus Glomus, the largest clade within the Glomeromycota. Higher AMF diversity was found in the least disturbed site, in which spores of Scutellospora persica, Glomus constrictum and Glomus globiferum were found in the rhizosphere of A. arenaria.

Plant-soil feedback as a mechanism of invasion by Carpobrotus edulis

Invasive plant species have been suggested to change the composition of the soil community in a way that results in a positive feedback for them and a negative feedback for the native plant community. Carpobrotus edulis, a species native to South Africa, is one of the most aggressive exotic species in Mediterranean Europe. Although several aspects of its invasion biology have been studied, the occurrence of plant-soil feedback has been scarcely investigated. We first checked for the existence of biotic resistance in soils from two invaded sites of Mediterranean Europe and one site in the native area. Secondly, we evaluated the effects of soil conditioning on the germination and plant growth of C. edulis and two key species of native dunes. Finally, we tested the effects of short- and long-term soil conditioning on the performance and reproductive effort of C. edulis. Our results show that at first there is a natural resistance to invasion by the soil biota. Later, biotic resistance in invaded soil is suppressed by the establishment of a soil community that enhances the growth of C. edulis and that negatively influences the growth and survival of the native plants. Long-term soil conditioning in the field resulted in shifts in the balance between vegetative growth and sexual reproduction. Long-term invasion was also reflected in high levels of endophyte colonization by chytrids in roots, although the physiological consequences of this colonization remain unknown. The results obtained illustrate a mechanism that explains how C. edulis breaks the initial biotic resistance of newly-invaded landscapes. Finally, this study highlights the importance of studying plant-soil interactions on different members of the plant community and temporal stages in order to fully understand invasion.

Genetic diversity and differentiation of Juniperus thurifera in Spain and Morocco as determined by SSR.

Juniperus thurifera L. is an important tree endemic to the western Mediterranean basin that it is able to grow in semi-arid climates. It nowadays exhibits a disjunct distribution pattern, occurring in North Africa, Spain, France and the Italian Alps. The Strait of Gibraltar has acted as an efficient barrier against gene flow between African and European populations, which are considered different subspecies by some authors. We aimed at describing the intraspecific genetic diversity of J. thurifera in populations from the Iberian Peninsula and Morocco and the phylogeographical relationships among these populations. The ploidy level of J. thurifera was examined and eleven nuclear microsatellites (nSSRs) developed for J. thurifera were assessed for genotyping this species. Six nSSRs were polymorphic and subsequently used to assess the genetic diversity and structure of the studied populations. Genotyping of the tetraploid J. thurifera using nuclear microsatellites supports the separation of Moroccan and Spanish populations into two genetically differentiated groups that correspond to the proposed subspecies africana and thurifera. High values of within population genetic diversity were found, that accounted for 90% of the total genetic variance, while population structure was weak. The estimators of genetic diversity were higher in populations of Spain than in populations of Morocco pointing for a possible loss of genetic diversity during the spread of this species to Africa from Europe.

A role for below‐ground biota in plant–plant facilitation

Summary 1. Plant–plant facilitation is an important driver of plant diversity, which in turn maintains ecosystem multifunctionality and can buffer some negative effects of climate change. Facilitation is classically attributed to the amelioration of environmental extremes and resource availability. Integrating below-ground biota into the positive plant interactions framework should provide a more realistic understanding of this process, enabling us to gain insights into the dynamics of below–above-ground communities. 2. We tested the effect of below-ground biota on the performance of a plant community and of individual species using soil extracts from the understorey of a benefactor plant species and adjacent open spaces. Soil bacteria from extracts and experimental microcosms were analysed using pyrosequencing. 3. Soil biota had a significant effect on the abundance, growth, functional traits and reproductive output of beneficiary plant species through processes that are independent of the direct influence of the benefactor species. Different soil bacterial communities were associated with the benefactor species, the individual beneficiary plant species and the plant community, revealing complex below– above-ground links between plants and soil microbiota. 4. Synthesis. The below-ground biota cultivated by benefactor plant species play a fundamental role in positive interactions between plant species contributing to the preservation of diversity and the evolution of plant communities.

Genetic Diversity and Differentiation of Ammophila arenaria (L.) Link as Revealed by ISSR Markers

Abstract Ammophila arenaria (L.) Link (marram grass) is the most important sand-fixing plant species along the northwestern European and Mediterranean coast, and it is also planted worldwide for sand dune stabilization. In spite of the intense use of this species in foredune restoration and stabilization programs, little is known about the genetic diversity within and between populations. We analyzed the genetic diversity of seven European populations of A. arenaria using inter-simple sequence repeats (ISSR) markers. The studied populations were selected in Wales, England, Belgium, the Netherlands, Portugal, and France. One half of the populations showed similar values of genetic diversity. The lowest values (Neis index =0.17) were found in the population from the Netherlands, which had been established after a foredune reinforcement project, and in a declining population in the south of Portugal. Statistical and phylo-genetic analyses revealed genetic differences between populations, and northern and southern clusters that corresponded to the two subspecies of A. arenaria. We propose that plant material for dune vegetation re-establishment programs should be collected locally rather than from remote populations that might be less well adapted to the local conditions of the planting site. However, when using local clonal plant material, care should be taken to collect the plant material from a number of sampling sites in order to ensure the genetic diversity of the new stands.

Genetic Diversity of Rhizobia Associated with Acacia longifolia in Two Stages of Invasion of Coastal Sand Dunes

ABSTRACT We examined the genetic diversity of root nodule bacteria associated with the Australian legume Acacia longifolia in two stages of invasion of a coastal sand dune system. All isolates belonged to the genus Bradyrhizobium. A higher diversity was found in the long-established trees. The results suggest the introduction of exotic bradyrhizobia with the plant.