Recovery of deep Posidonia oceanica meadows degraded by trawling

The recovery capacity of meadows of the Mediterranean seagrass Posidonia oceanica (L.) Delile in an area affected by illegal trawling were assessed after protection by anti-trawling reefs. The differences in vegetative growth between two impacted and two undisturbed localities were tested using growth, shoot balance, aborted branches, and leaf and rhizome production of both plagiotropic and orthotropic rhizomes. The organic matter in sediments, silt clay fraction and light intensity incident on the bottom were also measured in order to evaluate the physical conditions. Environmental and plant variables were measured in three sites placed inside each locality. The vegetative growth was positive in both impacted and control meadows but growth rates were lower in impacted than in control meadows. Average growth, production and shoot balance were greater in plagiotropic rhizomes from undisturbed localities (40.7±1.75 vs. 28.4±1.34 mm/year, 1133±0.06 vs. 708±0.04 mg DW/shoot/year, 1.36±0.08 vs. 0.96±0.06 shoots/year, respectively). Significantly greater values were also found in undisturbed localities for orthotropic rhizomes in terms of shoot balance and rhizome production (0.07±0.01 vs. 0.01±0.003 shoots/shoot/year and 155 vs. 124 mg DW/shoot/year, respectively). Of the physical parameters measured, only light intensity differed significantly between impacted and undisturbed localities. This parameter was 15.5% to 67.6% lower in impacted localities than in undisturbed localities, and this is the factor that causes the retardation of vegetative growth. The results show that recovery of P. oceanica meadows is possible after eliminating the cause of the impact. However, the very low rates of vegetative growth may prolong the time to total recuperation to almost 100 years. Therefore, effective management of P. oceanica meadows should aim to prevent meadow loss.     

A small-scale analysis of the spatial structure of a Posidonia oceanica meadow off the Island of Ischia (Gulf of Naples,Italy): Relationship with the seafloor morphology

Posidonia oceanica L. (Delile), an endemic species of the Mediterranean, forms extensive meadows which are continuously endangered by anthropic impacts. The availability of up-to-date information about interannual changes in shoot density of meadows and the knowledge of its expansion capabilities are crucial elements for the development of effective protection plans. Conversely, spatial ecology is becoming an increasingly important component of resource management, and the use of quantitative data for constructing prognosis maps of the dynamics of ecosystem degradation and restoration by nonlinear simulation methods is a topical field of landscape ecology. Unfortunately, little is known on spatial patterns of shoot density of P. oceanica on a small scale, despite their increasing use as indicators of the status and/or trends of meadows. The spatial structure of a continuous P. oceanica meadow, extending from 1 to 33 m depth in Lacco Ameno (Gulf of Naples, Italy), is investigated here by the “kriging” technique, a method widely used for geostatistical purposes. The analysis detected peculiar spatial patterns of shoot density and facilitated a small-scale (square meters) model of the distribution of P. oceanica. The highest shoot densities were found at the shallow stand (430 shoots m⁻², on the average, with a peak of 1000 shoots m⁻² in a relatively small area, at 1 m depth) and the lowest at the deep stand (average density <300 shoots m⁻² below 15 m depth). A high degree of patchiness was found in the shallow stand, down to 10 m depth. Nestlike patterns with a rounded shape, characterized by shoot density radially decreasing from the center, were demonstrated mainly in the shallow stand. An opposite trend was detected in one case, where the main nestlike pattern was characterized by a decrease of density towards the center. Nestlike patterns may be generated by the confluence and overlap of stolons expanding from proximal areas. The comparison of density and depth models indicated that the shape of the seafloor influences the density structure of meadows and the growth patterns of P. oceanica.     

Optimizing Posidonia oceanica (L.) Delile shoot density: Lessons learned from a shallow meadow

Although shoot patchiness has long been studied in Posidonia oceanica meadows, small scale spatial structure of P. oceanica meadows is poorly known, as very few studies focused on this feature. In order to analyze spatial patterns within P. oceanica meadows that appear uniformly dense and undisturbed, we collected shoot density data at the Capo Rizzuto Marine Protected Area (NE Mediterranean, Italy). Intensive sampling was carried out within a square lattice at small spatial scale (i.e. in the 10⁻¹–10² m² range) and shoot counts were obtained from sample quadrats of different size (60, 40, 20 cm). Spatial data analysis highlighted high irregularity in shoot density from centimetric to larger spatial scales. Therefore the deviation between shoot density estimates obtained using conventional methods and the overall average of quadrat counts (assumed as the best estimate for true density) was never negligible even when larger counting quadrats or higher numbers of replicates were adopted. While shoot density is regarded as the most important property of a P. oceanica meadow and as an indicator of ecosystem health, uncertainty in density estimates and unknown expected errors impair the effectivity of this approach. However, we showed that error could be predicted based on sampling intensity and design in an apparently uniform meadow. Although results from a single case study cannot be generalized, our work is the first attempt at analyzing the problems related to density estimates obtained from shoot counts and it shows how sampling can be optimized to achieve any desired level of accuracy.     

Effects of salinity on leaf growth and survival of the Mediterranean seagrass Posidonia oceanica (L.) Delile

The main aim of this study was to estimate the effects of salinity variation on the Mediterranean seagrass Posidonia oceanica (L.) Delile and its attached epiphytes. Leaf growth and survival of this plant were tested in several short-term (15 days) mesocosms experiments under controlled conditions between February 2001 and November 2001. Plants collected from shallow meadows at Alicante (SE Spain), with an ambient salinity of 36.8-38.0 psu, were placed in tanks of 300 L with an additional overhead light and exposed to different salinity treatments (ranging from 25 to 57 psu) during 15 days. To estimate the mortality and growth recuperation, in some experiments shoots were returned to control salinity (38 psu). Leaf growth was measured in the laboratory where epiphytic fauna and flora were removed from leaves, with a razor blade, to determine their biomass.P. oceanica was negatively influenced by increased salinity. Shoots showed a significant decrease in growth and survival, whereas epiphyte biomass did not show a clear response because of their high variability. Maximum leaf growth occurred between 25 and 39 psu. In addition, plants suffered considerable mortality at salinities above 42 psu and below 29 psu, with 100% mortality at 50 psu. In salinities between 39 and 46 psu, surviving plants were able to regain their original growth rate when returned to normal seawater salinity (38 psu). These results suggest that P. oceanica is one of the most sensitive seagrasses to salinity increments it is more tolerant to salinity reductions (25.0-36.4 psu), perhaps due to the terrestrial origin of seagrasses.     

Assessing effects of fishing prohibition on Posidonia oceanica seagrass meadows in the Marine Natural Reserve of Cerbère-Banyuls

The potential of alleviated fishing pressure measures established at the Marine Natural Reserve of Cerbère-Banyuls to affect phenological characters of P. oceanica, the main canopy-forming seagrass, was studied. Our results show differences in some leaf parameters between meadows under fishing prohibition measures compared to those without. In addition shallow P. oceanica meadows inside MPAs had lower non-structural carbohydrate content, yet are able to maintain themselves and, moreover, with an increased density. On the other hand, sexual reproduction was reduced inside protected areas.     

Seagrass light acclimation: 2-DE protein analysis in Posidonia leaves grown in chronic low light conditions

Posidonia oceanica meadows are among the most valuable coastal systems in the Mediterranean basin. They provide nursery and forage areas for many commercially important species, including juvenile mollusc, finfish, and crustaceans. In the Mediterranean Sea, P. oceanica beds have recently suffered from progressive die-offs attributed to lower light availability from elevated water turbidity. In order to understand adaptive low-light responses of this seagrass, we compared the protein expression in plants collected from turbid waters (low-light) with plants collected from pristine-clear waters (high-light). More than 2600 proteins were detected in leaves from both sites. Among them, 26 proteins were differentially expressed in low-light conditions, 12 of which were identified through MASCOT analyses. The remaining 14 proteins, did not receive significant identity scores due to a lack of genomic and proteomic information in available databases. Nevertheless, we observed a 30% down-regulation of RuBisCo large subunit in low-light acclimated leaves. Whereas, enzymes involved in carbohydrate cleavage (1-fructose-bisphosphate aldolase, nucleoside diphosphate kinase, and beta-amylase) were upregulated in low-light conditions. Electron microscopy studies also revealed substantial changes in the stroma lamellae/grana ratios in chloroplasts receiving low-light, possibly as a mechanism for re-establishing optimal PSI/PSII ratios. Furthermore, under low-light conditions, four components of the ubiquitin/mediated proteolysis pathway (26 S proteasome regulatory, proteasome beta type 1, proteasome 7 D beta type, and proteasome alpha 7), and the perchloric acid soluble translation inhibitor protein, were upregulated. This suggests that, in P. oceanica leaves, enhanced protein turnover mediates acclimation to low-light conditions. Also, enzymes involved in defending against cellular stress (superoxide dismutase, pyridoxine, and 2-caffeic-acido-methyl transferase) were differentially expressed in low-light regime. Subsequent aquaria studies involving P. oceanica transplants maintained in low- and high-light conditions, also demonstrate RuBisCo down-regulation and proteasomes upregulation in low-light acclimated plants.     

An improved method to monitor the health of seagrass meadows based on kriging

This paper focuses on the spatial pattern of the shoot density of the seagrass Posidonia oceanica through kriging methods. Kriging is a group of geostatistical techniques to predict the value of a field at an unobserved location from observations of its value at nearby locations. The results of an ordinary kriging of both raw values of shoot density and normalized shoot density with reference to a density vs. depth model (Dn) were compared. Results confirmed the existence of spatially organized parameters other than depth that influence patterns of P. oceanica shoot density. Although still requiring additional testing for complete validation, Dn mapping appears to be a valuable tool to (1) infer causes for the meadow's condition and to design a cost-effective field survey; (2) define areas where protection policies should be undertaken.     

Shoot age as a confounding factor on detecting the effect of human-induced disturbance on Posidonia oceanica growth performance

The response of orthotropic rhizome elongation and primary production of Posidonia oceanica to anthropogenic perturbations and potential confounding effects of shoot age were assessed using a Linear Multilevel Model (LMM). This model examined the confounding effect of age by comparing the estimates of impact and variance components obtained by excluding and including Age as an explanatory variable. Age had a negative effect on rhizome elongation and primary production with an annual decrease of 0.6 mm y^− 1 and 7 mg dw y^− 1 respectively. According to the LMM when age effect was omitted, the differences between disturbed and control locations in rhizome elongation and primary production were 2.62 mm y^− 1 and 0.044 g dw y^− 1 respectively. These effects were statistically not significant. On the contrary, when age effect was included in the statistical model, impacts became evident for both variables, with significant differences between disturbed and control locations of 5.85 mm y^− 1 and 0.081 g dw y^− 1 for rhizome elongation and primary production, respectively. Thus, particular attention should be paid to the potential confounding effect of shoots age in analyses of impacts on growth performance of P. oceanica.     

Anthesis effects on Posidonia oceanica (L.) Delile phenology in the Bay of Calvi (Corsica,Mediterranean Sea)

In Posidonia oceanica (L.) Delile, anthesis induces a decrease in the number of juvenile leaves resulting in a significant reduction in the number of leaves on the flowering shoots. All the leaves of the flowering shoots are narrower than the leaves of nonflowering shoots. A modification of the leaf growth also appears in flowering shoots: the oldest leaves are longer and the leaves induced during or after anthesis are shorter. At 10 m depth, in the Bay of Calvi, anthesis lasts roughly 3 months and the flowering is induced 7 months before anthesis.     

Protection Enhances Community and Habitat Stability: Evidence from a Mediterranean Marine Protected Area

Rare evidences support that Marine Protected Areas (MPAs) enhance the stability of marine habitats and assemblages. Based on nine years of observation (2001–2009) inside and outside a well managed MPA, we assessed the potential of conservation and management actions to modify patterns of spatial and/or temporal variability of Posidonia oceanica meadows, the lower midlittoral and the shallow infralittoral rock assemblages. Significant differences in both temporal variations and spatial patterns were observed between protected and unprotected locations. A lower temporal variability in the protected vs. unprotected assemblages was found in the shallow infralittoral, demonstrating that, at least at local scale, protection can enhance community stability. Macrobenthos with long-lived and relatively slow-growing invertebrates and structurally complex algal forms were homogeneously distributed in space and went through little fluctuations in time. In contrast, a mosaic of disturbed patches featured unprotected locations, with small-scale shifts from macroalgal stands to barrens, and harsh temporal variations between the two states. Opposite patterns of spatial and temporal variability were found for the midlittoral assemblages. Despite an overall clear pattern of seagrass regression through time, protected meadows showed a significantly higher shoot density than unprotected ones, suggesting a higher resistance to local human activities. Our results support the assumption that the exclusion/management of human activities within MPAs enhance the stability of the structural components of protected marine systems, reverting or arresting threat-induced trajectories of change.     

Spatial,temporal and structural variations of a Posidonia oceanica seagrass meadow facing human activities

The Bay of Saint-Cyr (Provence, France, Mediterranean Sea) is the site of two harbours, coastal urban development, trawling, boat anchoring and a sewage outfall. The Posidonia oceanica seagrass distribution was mapped with the help of aerial photographs, side scan sonar and GIS. In addition, the temporal variations of its distribution were studied by aerial photographs and GIS from 1955 to 2000. Finally, coverage and shoot density were measured via scuba-diving. This work reveals (i) the regression of the P. oceanica meadow at sites where harbours have been built, (ii) the occurrence of spaces within the meadow free of live P. oceanica (“intermattes”), which account for 8% of its surface area, (iii) a deep area where P. oceanica coverage and shoot density are low and (iv) evidence of regression, although modest, of the meadow at its lower limit. Nevertheless, the study site also exhibits an extensive and on the whole relatively healthy meadow whose limits have changed little over time.     

Increased death of new leaves of coastal banksia (Banksia integrifolia L.f) around ocean sewage outfall sites

Abstract Salt‐spray damage to coastal vegetation is a worldwide phenomenon. Worsening of such damage by surfactants present in sewage disposed of at sea has been observed in Europe and on trees planted around metropolitan beaches in Australia. Whether adverse impacts of sewage occur on the more widespread native vegetation around non‐metropolitan outfalls in Australia is unknown. The present study investigated the survivorship of existing leaves, and of new shoots and leaves, on Banksia integrifolia around three outfall and three non‐outfall sites in spring/summer of 1997‐1998. Survivorship of existing leaves was almost identical at outfall and non‐outfall sites. When plants initiated new shoots over the spring/summer, plants at outfall sites initiated more than plants at non‐outfall sites; but the proportion of these new shoots lost subsequently differed little between outfall and non‐outfall sites. Survivorship of newly initiated leaves on the new shoots differed between outfall and non‐outfall sites, with 65% of the new leaves on average being lost at outfall sites, and 47% being lost at non‐outfall sites by the end of summer. The probability of observing a difference of this magnitude ranged between P= 0.057 and 0.11 (February‐April 1998). This is the first report of a possible adverse impact on an Australian native plant from sewage discharged to the ocean.     

Marine space ecology and seagrasses. Does patch type matter in Posidonia oceanica seascapes?

The use of landscape tools in the study of seagrass meadows (seascapes) begins to be widely spread but still require the establishment of several basis, i.e. a patch type classification based on numerical characteristics. Thanks to the complex seascapes created by the Posidonia oceanica meadows, they appear to be suitable for a study at a patch type level (class), which bring a new insight of their arrangement at the whole seascape scale. By interpreting side scan sonar images from the Corsican coast (France) through a GIS software, it was possible to describe 11 types of patches and to evaluate their natural or anthropogenic origin. Comparison of different landscape metrics and wave exposure (Relative wave Exposure Index, REI) at the seascape and the patch level showed that the particularity of P. oceanica seascapes are mainly characterized by certain types of patches often of anthropogenic origin. Furthermore the REI seems not to be a relevant index for a study at a class scale. A bathymetrical succession of natural patches was outlined from the lower to the upper limit of the meadow, with a long-term dynamic opposed to a shorter one concerning anthropogenic patches. In order to assess the origin (natural or induced by human activities) of the patches in P. oceanica meadows, as well as in any other seagrass, a Patchiness Source Index (PaSI), ranging from 0 to 1, was defined.     

Boat anchoring on Posidonia oceanica beds in a marine protected area (Italy, western Mediterranean): effect of anchor types in different anchoring stages

Seagrasses worldwide are noted for suffering from mechanical damage caused by boat anchoring. This is particularly so in sites highly frequented by boaters (marine protected areas or coastal urbanised areas). In the last decades, different strategies have been put into practice to reduce such impacts on seagrasses (i.e. by anchoring bans or by deploying boat moorings). More recently, in consideration that few marine protected area (MPA) management bodies or local administrations have the resources to enforce their anchorage regulations, the self-regulatory approach based on education and information of boaters has been preferred in several cases. At present, however, very little is known on the correct anchoring practices to ensure the safeguarding of seagrasses. The aim of the present study was to experimentally quantify in the field the damage caused to Posidonia oceanica shoot density by anchoring. A multifactorial experiment was designed to test whether the damage is dependent on (1) different anchor types (Hall, Danforth and Folding grapnel), (2) the use of a chain vs. a rope, (3) the three anchoring stages (anchor fall, dragging/lock-in and weighing), and finally (4) whether the pattern is consistent among different locations of the meadow.As expected, the three anchor types employed in the present study differed in the levels of damage inflicted on the P. oceanica meadows of the Ustica Island MPA. In particular, the use of the Hall type anchor seems to be preferable to minimise this impact in comparison with the other two anchor types. Moreover, the effect on the meadow of the three anchor types is greatly dependent on the anchoring stage. These results confirm that the weighing stage is the critical stage of the anchoring process. The number of damaged shoots of P. oceanica was not affected by the presence of the chain. These patterns were consistent between locations.In the long term, even anchoring on P. oceanica by small boats using low-impact anchors may potentially have detrimental consequences. For this reason, we suggest that in vulnerable sites, it is preferable to implement an educational program based on information of boaters on correct anchoring practices and anchor typology to use, rather than adopting strong restrictions to boat anchoring or deploying mooring buoys. Although the use of these management strategies is still recommended in the case of anchorage frequented by bigger vessels using heavier anchors and chains.     

Leaf dimorphism in Thuja plicata and Platycladus orientalis (thujoid Cupressaceae s. str., Coniferales): the changes in morphology and anatomy from juvenile needle leaves to mature scale leaves

Thuja plicata and Platycladus orientalis initially produce only bifacial needle leaves. When the first lateral shoots develop, the leaf morphology and anatomy changes dramatically. Subsequently, only greatly reduced, bifacial scale leaves are developed. A new kind of “superimposed bifaciality” occurs with the change from juvenile needle leaves to mature scale leaves. Anatomical dorsiventrality affects not only the individual leaf, but also the complete plagiotropic lateral shoots of Thuja, which have a sun- and shade-exposed side. The upper light-exposed median leaves show adaxial leaf anatomy, contrary to the lower shaded median leaves showing abaxial leaf anatomy. Due to their mixed exposure, the lateral leaves show a lateral differentiation. At vertical lateral shoots of Platycladus, a predominant light-exposed side is absent. Thus, the anatomical dorsiventrality does not affect the complete shoot. Here the morphological abaxial side of a scale leaf becomes functionally and physiologically adaxial by reorientation of the palisade parenchyma and stomata. In juvenile needle leaves, the palisade parenchyma is located adaxial, with the majority of stomata being located abaxial. Conversely, in mature scale leaves, the palisade parenchyma is abaxial and the majority of stomata are adaxial.     

Vicariance patterns in the Mediterranean Sea: east–west cleavage and low dispersal in the endemic seagrass Posidonia oceanica

Aim The seagrass, Posidonia oceanica is a clonal angiosperm endemic to the Mediterranean Sea. Previous studies have suggested that clonal growth is far greater than sexual recruitment and thus leads to low clonal diversity within meadows. However, recently developed microsatellite markers indicate that there are many different genotypes, and therefore many distinct clones present. The low resolution of markers used in the past limited our ability to estimate clonality and assess the individual level. New high‐resolution dinucleotide microsatellites now allow genetically distinct individuals to be identified, enabling more reliable estimation of population genetic parameters across the Mediterranean Basin. We investigated the biogeography and dispersal of P. oceanica at various spatial scales in order to assess the influence of different evolutionary factors shaping the distribution of genetic diversity in this species. Location The Mediterranean. Methods We used seven hypervariable microsatellite markers, in addition to the five previously existing markers, to describe the spatial distribution of genetic variability in 34 meadows spread throughout the Mediterranean, on the basis of an average of 35.6 (± 6.3) ramets sampled. Results At the scale of the Mediterranean Sea as a whole, a strong east–west cleavage was detected (amova) . These results are in line with those obtained using previous markers. The new results showed the presence of a putative secondary contact zone at the Siculo‐Tunisian Strait, which exhibited high allelic richness and shared alleles absent from the eastern and western basins. F statistics (pairwise θ ranges between 0.09 and 0.71) revealed high genetic structure between meadows, both at a small scale (about 2 to 200 km) and at a medium scale within the eastern and western basins, independent of geographical distance. At the intrameadow scale, significant spatial autocorrelation in six out of 15 locations revealed that dispersal can be restricted to the scale of a few metres. Main conclusions A stochastic pattern of effective migration due to low population size, turnover and seed survival is the most likely explanation for this pattern of highly restricted gene flow, despite the importance of an a priori seed dispersal potential. The east–west cleavage probably represents the outline of vicariance caused by the last Pleistocene ice age and maintained to this day by low gene flow. These results emphasize the diversity of evolutionary processes shaping the genetic structure at different spatial scales.     

The role of disturbance in promoting the spread of the invasive seaweed Caulerpa racemosa in seagrass meadows

Human disturbances, such as anchoring and dredging, can cause physical removal of seagrass rhizomes and shoots, leading to the fragmentation of meadows. The introduced green alga, Caulerpa racemosa, is widely spread in the North-West Mediterranean and, although it can establish in both degraded and pristine environments, its ability to become a dominant component of macroalgal assemblages seems greater in the former. The aim of this study was to estimate whether the spread of C. racemosa depends on the intensity of disturbance to the canopy structure of Posidonia oceanica. A field experiment was started in July 2010 when habitat complexity of a P. oceanica meadow was manipulated to simulate mechanical disturbances of different intensity: rhizome damage (High disturbance intensity = H), leaf removal (Low disturbance intensity = L), and undisturbed (Control = C). Disturbance was applied within plots of different size (40 × 40 cm and 80 × 80 cm), both inside and at the edge of the P. oceanica meadow, according to an orthogonal multifactorial design. In November 2011 (16 months after the start of the experiment), no C. racemosa was found inside the seagrass meadow, while, at the edge, the cover of the seaweed was dependent on disturbance intensity, being greater where the rhizomes had been damaged (H) than in leaf removal (L) or undisturbed (C) plots. The results of this study indicate that physical disturbance at the margin of seagrass meadows can promote the spread of C. racemosa.     

Genetic Polymorphism and Transplantation Success in the Mediterranean Seagrass Posidonia oceanica

We studied the role of genetic variability of donor beds in establishing transplantation criteria for the Mediterranean seagrass Posidonia oceanica. Horizontal rhizomes, collected from three geographically distinct populations, were transplanted into a common bed at a highly human‐impacted locality. The transplantation site was located near one of the donor populations. After three years, the shoots collected in the population adjacent to the transplanting site showed the lowest growth performance. Genetic variability, assessed through the analysis of hypervariable microsatellite regions, and growth performance followed a similar trend. The shoots growing and branching at the highest rate were those collected from populations with the highest heterozygosity values, despite greater geographic distance. No genetic differences were found between the transplanted shoots and shoots from donor meadows, as expected due to the low rate of sexual reproduction in P. oceanica and the short time that had passed since the transplants. The problem of affecting the local gene pool by the introduction of foreign genotypes could arise, but introduction of new alleles could balance the degradation of genetic variability caused by human impact. In general our study suggests that the genetic variability of source material is an important aspect to consider in the development of seagrass restoration strategies.     

Against the odds: complete outcrossing in a monoecious clonal seagrass Posidonia australis (Posidoniaceae)

Background and Aims

Seagrasses are marine, flowering plants with a hydrophilous pollination strategy. In these plants, successful mating requires dispersal of filamentous pollen grains through the water column to receptive stigmas. Approximately 40 % of seagrass species are monoecious, and therefore little pollen movement is required if inbreeding is tolerated. Outcrossing in these species is further impacted by clonality, which is variable, but can be extensive in large, dense meadows. Despite this, little is known about the interaction between clonal structure, genetic diversity and mating systems in hydrophilous taxa.

Methods

Polymorphic microsatellite DNA markers were used to characterize genetic diversity, clonal structure, mating system and realized pollen dispersal in two meadows of the temperate, monoecious seagrass, Posidonia australis, in Cockburn Sound, Western Australia.

Key Results

Within the two sampled meadows, genetic diversity was moderate among the maternal shoots (R = 0·45 and 0·64) and extremely high in the embryos (R = 0·93–0·97). Both meadows exhibited a highly clumping (or phalanx) structure among clones, with spatial autocorrelation analysis showing significant genetic structure among shoots and embryos up to 10–15 m. Outcrossing rates were not significantly different from one. Pollen dispersal distances inferred from paternity assignment averaged 30·8 and 26·8 m, which was larger than the mean clone size (12·8 and 13·8 m).

Conclusions

These results suggest highly effective movement of pollen in the water column. Despite strong clonal structure and moderate genetic diversity within meadows, hydrophilous pollination is an effective vector for completely outcrossed offspring. The different localized water conditions at each site (highly exposed conditions vs. weak directional flow) appear to have little influence on the success and pattern of successful pollination in the two meadows.