Patterns of seagrass (Posidonia oceanica) flowering in the Western Mediterranean

The intensive reconstructive sampling (1957–2004, 39 localities), a systematic direct observation (1992–2004, 1 locality) and particular direct observations (66 localities) of Posidonia oceanica meadows were analysed together with temporal series of flowering available in the literature (19 localities). This allowed the examination of temporal and spatial variability in annual flowering prevalence (FP, the fraction of meadows flowering in a given year) and of annual meadow flowering intensity (FI, number of inflorescences per shoot) for the period 1979–2004 across the Western Mediterranean, as well as spatial variability of flowering frequency (FF, the fraction of years that a given meadow has flowered) and shoot flowering probability (Pf, fraction of flowering stalks appeared per annual segment). Each year, on an average 17% of the investigated meadows flowered, ranging from 3 to 86% of meadows among the years. The highest annual FP and FI values were obtained in 2003 (FP=0.86 and mean FI=0.23±0.03 inflorescences shoot⁻¹). A secondary peak of FP and mean FI occurred 9 years earlier, in 1994 (FP=0.44 and mean FI=0.08±0.02). Both peaks of flowering occurred after hot summers. Flowering synchrony in particular years across the Western Mediterranean and clines of increased meadow flowering frequency towards the North and East, suggests the existence of large-scale environmental mechanisms controlling the floral induction. On the other hand, meadow FF and Pf were highly heterogeneous among and within the meadows, indicating that local factors also may play a significant role in flowering induction. When flowering, the Western Mediterranean meadows showed an average 0.11±0.02 inflorescences shoot⁻¹, but FI greatly varied among and along the series (from 0.002 to 0.54 inflorescences shoot⁻¹) and decreased significantly with depth but was independent of meadow shoot density and meadow latitude or longitude. The shoot flowering probability was quite low (0.007±0.002 inflorescences shoot⁻¹ year⁻¹) and exponentially increased with shoot age.     

Primary production of Posidonia oceanica in the Mediterranean Basin

Primary production of the marine phanerogam Posidonia oceanica (Linnaeus) Delile was measured by lepidochronological analyses at 22 sites in the Mediterranean Sea (Corsica, France, Italy, Sardinia and Turkey), between 1983 and 1992, to determine spatial and temporal variations. Leaf production (blade and sheath) ranged from 310 to 1 540 mg dry wt shoot^–1 yr^–1, depending on site and depth. Rhizome production ranged from 24 to 120 mg dry wt shoot^–1 yr^–1 (6% of average leaf production). At some sites the results obtained by lepidochronological analysis were consistent with earlier results obtained by classic methods (e.g. leaf-marking). While primary production per shoot (mg dry wt shoot^–1 yr^–1) displayed no significant differences between sites, primary production of the P. oceanica meadow (g dry wt m^–2 yr^–1) decreased with increasing depth at all sites studied. This decrease correlated with reduced density of the meadow (number of shoots per m²) with increasing depth. Past primary production was also extrapolated at three sites at the island of Ischia (Italy) for a period of 5 yr in order to determine interannual variations over a period of several years. While major variations were recorded for the surface stations (5 and 10 m depth), production remained stable at the deepest station (20 m depth). Given the large geographical scale of the study (location, depth range), it would appear that while P. oceanica production remains considerable, the values recorded in the literature on the basis of classical analyses (surface stations) represent maxima, and cannot be generalised for meadows as a whole.     

Characterization of the benthic vegetation in the Farwà Lagoon (Libya)

This study forms part of the Action Plan for the Conservation of Marine Vegetation in the Mediterranean SEa (United Nation Environmental Program). It was carried out in June 2000 in the Farwà Lagoon, Libya. The mapping of the main benthic vegetation was achieved by compiling the field observations (transect method), and remote sensing of SPOT satellite images. The phytobenthos in the Farwà lagoon covers an area of 1820 ha (65%). Three benthic macrophyte species dominate, namely the marine phanerogamsCymodocea nodosa andPosidonia oceanica, and the algaCaulerpa prolifera. DeadPosidonia oceanica leaves (litter) form veritable mounds in the vicinity of the openings leading to open sea. These leaves, which come from the coastal sea, are brought into the lagoon by currents and tides; their decomposition will lead to high oxygen consumption and the release of hydrogen sulphide. The phenological data ofPosidonia oceanica shoots sampled in the lagoon are similar to those from other stations in the Mediterranean. Conversely, the lepidochronological parameters of shoots sampled in the central part of the lagoon exhibit values that are substantially higher than those generally recorded in the Mediterranean. The mean number of leaves produced annually is 9.9 (mean value for the Mediterranean: 7.5) and the rhizome growth rate is of 35.7 mm.yr⁻¹ (mean value for the Mediterranean: 7.5 mm.yr⁻¹). This hypersaline environment would seem to provide optimum growth conditions for the speciesPosidonia oceanica.     

Mixed but not admixed: a spatial analysis of genetic variation of an invasive ascidian on natural and artificial substrates

Following the introduction to a new area (pre-border dispersal), post-border processes determine the success in the establishment of non-indigenous species (NIS). However, little is known on how these post-border processes shape the genetic composition of NIS at regional scales. Here, we analyse genetic variation in introduced populations along impacted coastlines to infer demographic and kinship dynamics at the post-border stage. We used as a model system the ascidian species Microcosmus squamiger that has been introduced worldwide. This species can colonize and grow fast on man-made artificial structures, impacting activities such as mariculture. However, it can also establish itself on natural substrates, thus altering natural communities and becoming an ecological problem. We genotyped 302 individuals from eight populations established on natural and artificial substrates in the north-western Mediterranean Sea, using six microsatellite loci. We then compared the resulting genotypes with those found within the native range of the species. We found high levels of genetic diversity and allelic richness in all populations, with an overall deficit of heterozygotes. Autocorrelation analyses showed that there was no within-population genetic structure (at a scale of tens of metres); likewise, no significant differentiation in pairwise comparisons between populations (tens of kilometres apart) and no isolation-by-distance pattern was found. The results suggest that M. squamiger has a natural capacity for high dispersal from one patch of hard substrate to another and no differences whatsoever could be substantiated between natural and artificial substrates. Interestingly, two groups of genetically differentiated individuals were detected that were associated with the two ancestral source areas of the worldwide expansion of the species. Individual assignment tests showed the coexistence of individuals of these two clusters in all populations but with little interbreeding among them as the frequency of admixed individuals was only 15 %. The mechanism responsible for maintaining these genetic pools unmixed is unknown, but it does not appear to compromise post-border colonization of introduced populations.     

Wanted dead or alive: high diversity of macroinvertebrates associated with living and ‘dead’ Posidonia oceanica matte

The Mediterranean endemic seagrass Posidonia oceanica forms beds characterised by a dense leaf canopy and a thick root-rhizome ‘matte’. Death of P. oceanica shoots leads to exposure of the underlying matte, which can persist for many years, and is termed ‘dead’ matte. Traditionally, dead matte has been regarded as a degraded habitat. To test whether this assumption was true, the motile macroinvertebrates of adjacent living (with shoots) and dead (without shoots) matte of P. oceanica were sampled in four different plots located at the same depth (5–6 m) in Mellieha Bay, Malta (central Mediterranean). The total number of species and abundance were significantly higher (ANOVA; P<0.05 and P<0.01, respectively) in the dead matte than in living P. oceanica matte, despite the presence of the foliar canopy in the latter. Multivariate analysis (MDS) clearly showed two main groups of assemblages, corresponding to the two matte types. The amphipods Leptocheirus guttatus and Maera grossimana, and the polychaete Nereis rava contributed most to the dissimilarity between the two different matte types. Several unique properties of the dead matte contributing to the unexpected higher number of species and abundance of motile macroinvertebrates associated with this habitat are discussed. The findings have important implications for the conservation of bare P. oceanica matte, which has been generally viewed as a habitat of low ecological value.     

Animals on marine flowers: does the presence of flowering shoots affect mobile epifaunal assemblage in an eelgrass meadow?

Eelgrass, Zostera marina, produces two types of shoots: morphologically simple vegetative shoots and highly branched flowering (reproductive) shoots, the latter found only in summer months. We examined whether the abundance and diversity of mobile epifaunal assemblage are affected by the presence of flowering shoots in an eelgrass meadow of Otsuchi Bay, northeastern Japan. Comparisons of epifauna in natural vegetation revealed that density and species richness did not differ significantly between sites consisting of both flowering and vegetative shoots, and those only of vegetative shoots. A transplant experiment, conducted to examine the colonization rates of epifauna to defaunated eelgrass planted with different combination of vegetative and flowering shoots, showed no obvious variation in abundance and species richness. At species level, the density of some species such as a tanaid Zeuxo sp. and a polychaete Platynereis sp. was higher at sites and/or treatments with flowering shoots, whereas that of some gastropods, such as Lirularia iridescens and Siphonacmea oblongata was higher at sites without flowering shoots. The species-specific response led to dissimilarity of epifaunal assemblage between sites and among treatments with different densities of vegetative and flowering shoots. Similar patterns observed for natural vegetation and the transplant experiment suggest that the variation in assemblage structure is caused by habitat selection of each species, for example, the utilization of flowering shoots as feeding ground and nursery by Zeuxo sp.     

Supply-side ecology of the brown mussel,<Emphasis Type="Italic"> Perna perna</Emphasis>: an investigation of spatial and temporal variation in,and coupling between,gamete release and larval supply

Sampling of recruitment-associated variables of Perna perna was done approximately monthly for 14 months at intertidal locations 500 m apart, nested within sites 25 km apart. Paired with intertidal locations were nearshore locations, 600 m to sea. Sampling assessed spawning, densities of larvae in the water column and densities of late plantigrades and juveniles on the shore. Major events in each variable were synchronous over larger scales (10s of kilometres) while subsidiary events were synchronised at smaller scales, varying within sites (100s of metres) or even within locations (metres). This suggests that the processes driving major events operated over large scales while processes operating at much more local scales drove less intense, more localised events. A major spawning event occurred at all locations in May–June 1998. Weaker spawning events occurred at different times in different locations. Larvae were found on 80% of sampling occasions, densities peaking in January–March 1998 and 1999 at all locations. Plantigrades and juveniles showed less clear patterns, with considerable residual variation. There was no sign of strong coupling among variables with few significant direct or cross correlations. The major sources of variability shifted from time to space as one progressed from spawning, to plantigrade density to juvenile density. For spawning, time was the most important source (58%) of heterogeneity and space accounted for little (8%) of the total variance. For larvae and late plantigrades, time was still the most important source of variability (41% and 33%, respectively), but space was a much more substantial component. For juveniles, small-scale (residual) spatial variability dominated total variability (75%). This strongly suggests the importance of hydrography and its effects on variation in delivery of larvae to the intertidal from offshore. These findings also indicate greater spatial heterogeneity as recruits age, reflecting small-scale variations in larval delivery and the increasing importance of post-settlement mortality.Communicated by G.F. Humphrey, Sydney     

Environmental determinants of distribution and foraging behaviour of cormorants (Phalacrocorax spp.) in temperate estuarine habitats

The distribution and behaviour of cormorants in estuarine environments was examined on the central coast of New South Wales, Australia, with respect to habitat associations at different spatial scales. No consistent variation in abundance was found for four species of cormorants (great Phalacrocorax carbo, pied P. varius, little black P. sulcirostris, and little pied P. melanoleucos) with state of tide (high and low) and time of day (early, middle, and late) at five estuarine locations. Differences in abundance were found among locations that were not confounded by short-term temporal variation (i.e. time of day and tide). Differences in abundance were detected among habitats (e.g. bays, creeks, and headlands) separated by hundreds of metres to kilometres in different estuaries. Cormorants of all species were rare on the open coast and near the entrance of estuaries. Abundances of cormorants varied greatly within and among creeks, bays, and river channels. Presence of seagrass beds explained much of this variation and most of the cormorants swimming and feeding were found near seagrass. Mapping of seagrass beds and the positions of cormorants at scales of metres to hundreds of metres showed a close relationship between the presence of swimming and roosting beds and the presence of seagrass beds for P. melanoleucos and for P. carbo. We argue that cormorants make decisions to visit particular estuarine habitats, especially those with seagrass, where many types of prey (e.g. fish and crustaceans) are probably most abundant. These choices must be interpreted in the context of decisions that cormorants make on scales of hundreds to thousands of kilometres during periodic excursions to the interior of Australia. Further, environmental threats to seagrass beds could impinge on these mobile visitors to the same extent as on more permanent residents. Received: 14 February 2000 / Accepted: 17 July 2000     

Genetic differentiation among populations of the Mediterranean hermaphroditic brooding coral<Emphasis Type="Italic"> Balanophyllia europaea</Emphasis> (Scleractinia: Dendrophylliidae)

Spatial models of genetic structure and potential gene flow were determined for five populations of Balanophyllia europaea, a simultaneous hermaphroditic and brooding coral, endemic to the Mediterranean. Six allozyme loci indicated a genetic structure that departed markedly from Hardy–Weinberg equilibrium, with a significant lack of heterozygotes. The genetic structure observed supports the hypothesis that self-fertilisation characterises the reproductive biology of B. europaea. Populations at small spatial scales (8–40 m) are genetically connected, while those at large scales (36–1,941 km) are genetically fragmented; the genetic differentiation of the populations is not correlated to geographic separation. This spatial model of genetic structure is compatible with an inbreeding mating system. Furthermore, it is also consistent with the expected dispersal potential of swimming larvae of brooding corals, i.e. larvae that are able to produce significant gene flows only within limited spatial scales.Communicated by R. Cattaneo-Vietti, Genova     

Reproductive synchrony in a habitat-forming kelp and its relationship with environmental conditions

Key biological processes such as the timing and synchrony of reproduction, are sensitive to fluctuations in the environment, as well as alterations in the physiology and behaviour of individuals and the ecology of populations. The main aim of this study was to identify patterns in reproductive timing and relationships with prevailing environmental conditions, for the major habitat-forming alga in temperate Australia, Ecklonia radiata (C. Agardh) J. Agardh. We observed strong synchronous patterns of zoospore production across spatial scales from hundreds of metres to tens of kilometres, with populations of E. radiata reproducing from mid-summer to the end of autumn. High zoospore densities were positively related to sea temperature and E. radiata released zoospores in temperatures well above previously documented thermal tolerance limits for other Laminariales. Reproductive timing was also negatively related to seasonal increase in wave exposure. We conclude that E. radiata undergoes synchronous reproduction which can be linked to its annual growth cycle and natural environmental fluctuations, to promote growth and survival of recruits.     

Polychaetes associated to a<Emphasis Type="Italic"> Cymodocea nodosa</Emphasis> meadow in the Canary Islands: assemblage structure,temporal variability and vertical distribution compared to other Mediterranean seagrass meadows

The structure, diversity and temporal distribution of the infaunal polychaetes associated with Cymodocea nodosa meadows were studied in Tenerife (Canary Islands). The samples were collected monthly throughout a year, to depths of 13–16 m. The sediment was extracted by means of PVC cores, in which four layers were separated (i.e. 0–5 cm, 5–10 cm, 10–20 cm and 20–30 cm). A total of 1,167 polychaete specimens, belonging to 69 taxa were collected, representing one of the most dominant groups in the benthic assemblage throughout the entire year. The most common families were Syllidae, Paraonidae and Spionidae, both in terms of abundance and species richness. The dominant species were Streptosyllis bidentata, Aricidea assimilis and Exogone parahomoseta mediterranea, representing also the only constant species throughout the year. The highest values of species richness, diversity, equitability and abundance of polychaetes occurred in September. The multifactorial analysis of abundances (i.e. cluster analysis and non-metric, multi-dimensional scaling) indicated temporal segregation of the samples from July, August and September (i.e. the warmest months) with respect to those from the rest of the year, due to structural differences in the assemblage. Polychaete species have been found to a depth of up to 30 cm in the sediment. Nevertheless, most of them (89%) occurred in the upper 5 cm of the sediment, with an increase of specimens in deeper layers in February (i.e. due to sporadic episodes of higher hydrodynamics). To compare the vertical distribution of polychaetes, additional core samples were collected in two seagrass meadows (i.e. C. nodosa and Ruppia cirrhosa) at Ebros Delta (NW Mediterranean); these were separated into five layers (i.e. 0–5 cm, 5–10 cm, 10–15 cm, 15–20 cm, 20–25 cm). The results obtained for the R. cirrhosa meadow (98% of the polychaetes within the upper 5 cm) agree with those for the Canarian C. nodosa meadow, while the polychaetes reached up to 15 cm depth in the Mediterranean C. nodosa meadow (i.e. ~39% between 0 and 5 cm, ~41% between 5 and 10 cm, ~20% between 10 and 15 cm). Our results indicated that the structural characteristics of the assemblages appeared to be more strongly controlled by the combined characteristics of the sediment (i.e. lack of oxygen, granulometry and degree of compaction) than by the seagrass species building the meadow.Communicated by O. Kinne, Oldendorf/Luhe     

Reproduction in the seagrass Zostera novazelandica on intertidal platforms in southern New Zealand

This study investigates the reproductive periodicity and reproductive output of the seagrass Zostera novazelandica on two intertidal reefs. Peak numbers of flowering shoots occurred during March (late summer) of two years at both sites and no flowering shoots occurred during the winter months of July to September. There were greater numbers of flowering shoots in seagrass patches in the low intertidal zone (up to an average of 55 per 0.1 m²) compared to the middle (up to 20 per 0.1 m²) and upper (up to 9 per 0.1 m²) zones, and about three times greater reproductive output in patches associated with tidepools compared to those not bordering tidepools. The average number of inflorescences per shoot was 3.1 (±0.25) at one site vs 1.2 (±0.08) at the other, and showed a progressive decrease from the lower shore to the upper shore. Patches associated with tidepools had twice the number of inflorescences per shoot (2.8 ± 0.24) than patches not bordering tidepools (1.5 ± 0.16). The number of flowering shoots was highly correlated with leaf width, leaf length, and ramet density, while the leaf-area index decreased from the lower shore to the upper shore. The reproductive effort of plants, as measured by the percent biomass invested in flowering shoots during peak reproduction, was significantly different between sites, tidepool associations, and shore level. For all the variables measured, there was considerable spatial variation, with significant interaction terms between most factors investigated. In laboratory experiments, more inflorescences were produced at light intensities of 30 and 300 μE m⁻² s⁻¹ than at 100 μE m⁻² s⁻¹. At a salinity of 17‰, 1.5 × the number of flowers was produced than at 33‰, while none was formed at 70‰. Plants cultured at 5 °C had about three times the number of inflorescences than those at 15 °C, while none was formed at 25 °C. Received: 25 June 1997 / Accepted: 24 September 1997     

Seagrass mortality due to oversedimentation: an experimental approach

Mortality due to oversedimentation of the Mediterranean seagrassPosidonia oceanica was experimentally evaluated by field manipulations of the sediment level. Increased levels of sediment placed over plant shoots and rhizomes induced significant shoot mortality, even at moderate burial levels (ca. 5 cm). When sediment was added to reach levels 15 cm higher than the initial one, a 100 % mortality was observed after 200–300 days. The response of the plant was independent of site and depth. These results can be used in ecological risk assessment of coastal activities which potentially affect sediment deposition.     

Chemistry,mineralogy and radioactivity in posidonia oceanica meadows from North-Western Sicily

This paper presents measurements of major, minor, metal trace elements and radionuclides in sediments and in Posidonia oceanica samples from north-western Sicily (Italy). The mineralogical and chemical composition of sediments were determined by X-ray diffraction and X-ray fluorescence techniques, respectively. A flame atomic absorption spectrophotometry was used to measure concentrations of Cu, Zn, Cd and Pb in P. oceanica samples and in sediments. Specific activities of selected radionuclides have been determined by high-resolution gamma spectrometry. Standard statistical analysis was used to assess correlations between different elements and different sample types.     

Distribution patterns of sea urchins and barrens in shallow Mediterranean rocky reefs impacted by the illegal fishery of the rock-boring mollusc<Emphasis Type="Italic"> Lithophaga lithophaga</Emphasis>

Shallow rocky habitats in SW Apulia (SE Italy, Mediterranean Sea) were surveyed in late spring 2002 to assess distribution patterns of sea urchins (Paracentrotus lividus and Arbacia lixula) and barren habitats (coralline barrens and bare substrates) in rocky reefs impacted by the destructive fishery of the rock-boring date-mussel Lithophaga lithophaga. Sea urchin density, test size-structure and biomass, and the percent cover of barrens were evaluated at four locations (5–6 km apart from each other), two heavily impacted by the date-mussel fishery and two controls. Sea urchin density and barren habitat cover were assessed at two and three sites (100–300 m apart), respectively, within each location. Sea urchin biomass was evaluated only at the scale of locations. Average density of P. lividus did not significantly change between impacted locations and controls, whereas A. lixula showed a greater density at the impacted locations. Distribution patterns of A. lixula, in addition, differed at the spatial scale of a few metres between impacted locations and controls, being generally more aggregated at the controls. The size-frequency distribution (test diameter) of P. lividus showed a mode at 3–4 cm at the impacted locations compared to a mode at 2–3 cm in the controls. The size-frequency of A. lixula was bimodal at the damaged locations (with modes at 1–2 and 4–5 cm, respectively) and unimodal (with the mode at 4–5 cm) at the controls. Average biomass of both sea urchins (P. lividus and A. lixula) was two- to fourfold greater at the impacted locations (~600 g wet wt m^–2) than at the controls (150–250 g wet wt m^–2). Barren habitats had a far greater average cover (mainly of macroalgae) at the impacted locations (from 79% to 96%) than at control locations (from 7% to 21%). These results show that the date-mussel fishery may have the potential to affect distribution patterns of sea urchins and to greatly enhance the percent cover of barren grounds in shallow Mediterranean rocky reefs.Communicated by R. Cattaneo-Vietti, Genova     

Geographic,population, and seasonal patterns in the reproductive parameters of the sea urchin <Emphasis Type="Italic">Paracentrotus lividus</Emphasis>

Generalized additive models were used to synthesize the data from 52 publications on the gonad growth and spawning seasonality of Paracentrotus lividus and identify spatial and temporal patterns in the reproductive processes. According to our results, Atlantic populations spawn in spring and develop large gonads (6.4 ± 0.2% of fresh body weight (mean ± SE)), which increase in size toward higher latitudes. While in the Mediterranean, the gonads are smaller (3.1 ± 0.1%), possibly as the result of successive spawnings which impede nutrients accumulating in the gonads. Differences were also observed between habitats: gonad production was higher in seagrass meadows in the Mediterranean and in subtidal rocky substrates in the Atlantic. Food availability might cause these variations as well as the negative correlation observed between depth and the gonad index for rocky substrates. Sex and body size do not seem to influence greatly gonad growth.     

Variations in mortality of a coral-reef fish: links with predator abundance

The mortality rates of a pomacentrid Acanthochromis polyacanthus were examined in relation to the abundance of large predatory fish (>200 mm total length, TL) at two spatial scales. Survivorship was negatively related to patterns of predator abundance at a large spatial scale (hundreds of metres) over 3 yr, but not at a small spatial scale (tens of metres) over 2 yr. On the large scale, mortality was consistently greater (14 to 30%) in locations where there were greater numbers of predators, and lower in locations where predators occurred in smaller numbers. Among these locations, spatial differences in rank abundance of surviving juveniles were primarily due to mortality, whereas temporal differences in rank abundance were primarily due to initial juvenile abundance. These data suggest that impacts of large predatory fish were likely to have been greater in space than time and at the large spatial scale than the small spatial scale.     

Patterns in reproductive dynamics of burrowing ghost shrimp Trypaea australiensis from small to intermediate scales

Many studies have examined latitudinal differences in reproduction of marine invertebrates, but few have measured variation at small to intermediate scales (kilometres to hundreds of kilometres), which may confound comparisons across broader geographic regions. Here, we examined variation in the reproductive biology of a little-studied species of burrowing ghost shrimp (Trypaea australiensis) at spatial scales ranging from km (between sites within estuaries) to 100s of km (among estuaries), over a 2-year period in south-eastern Australia. Sex ratios of populations were consistently biased towards females through time and space. Although reproduction started in summer months across all spatial scales, there was a pattern of earlier spawning from southern to northern estuaries. Integration of results from previous studies of T. australiensis supported a similar pattern of earlier breeding from high to low latitudes. Fecundity of shrimp increased linearly with female size, but the relationship varied inconsistently across the different spatial scales. Similarly, sizes at maturity varied from small to intermediate scales and observed patterns were not consistent with general predictions e.g. shrimp were smaller and ovigerous at smaller sizes at sites in the southern-most estuary, compared to estuaries further north. We found no differences in the sizes of embryos across the different spatial scales, but confirm that T. australiensis employs a strategy of high fecundity and small embryo size compared to other thalassinidean shrimp. Our results suggest that factors at smaller scales (e.g. food availability) may be important in affecting reproductive dynamics of T. australiensis, but further research is needed in testing hypotheses about patterns observed here. A lack of similar studies on other marine organisms remains an impediment to understanding life-history strategies and the sustainable management and conservation of populations.     

Population age structure and rhizome growth of<Emphasis Type="Italic"> Cymodocea nodosa</Emphasis> in the Ria Formosa (southern Portugal)

The population age structure and derived population dynamics (recruitment, growth and mortality), rhizome growth and flowering effort of Cymodocea nodosa in the Ria Formosa (south of Portugal) were examined using reconstruction techniques. The horizontal rhizome elongation rates were low (13.8–30.7 cm year^–1), which is consistent with the low population growth rate (–2.35 to 0.29 year^–1). The vertical elongation rate was proportional to the horizontal growth except for one stand that was subjected to sand movements and had higher elongation rates. The mean leaf production varied from 11.8 to 14 leaves year^–1. The average shoot age varied from 0.70 to 1.04 years, and the oldest shoot found was 7.6 years. The mortality rates were the highest as yet reported for this species (0.99–3.70 year^–1). Observed low growth and high mortality are possibly associated with the development of eutrophic conditions in the Ria Formosa. The flowering probability found in Ria Formosa stands (<0.009–0.90 shoots^–1 year^–1) was comparable to that found in the Mediterranean and off the Canary Islands. Only two stands with flowering events were found inthe Ria Formosa population, which indicates that the stands are highly dependent on clonal growth to maintain themselves inthe Ria Formosa.Communicated by O. Kinne, Oldendorf/Luhe