Detrital stocks and dynamics of the seagrass Posidonia oceanica (L.) Delile in the Spanish Mediterranean

Previous studies have shown that most leaf production (>90%) of the seagrass Posidonia oceanica is shed after senescence and that a substantial percentage (up to 80%) may thereafter be exported off the seagrass meadows by waves and currents. It has also been reported that P. oceanica meadows can accumulate large stocks of belowground detritus due to slow decomposition rates. However, the generality of these results across broad spatial scales is poorly known. In this report, we examine the fate of leaf production and the magnitude and dynamics of belowground detritus in 16 P. oceanica meadows distributed along the Spanish Mediterranean. Herbivores removed a small percentage of leaf production in all the meadows (≤13%), with most leaf production (>85%) being shed after senescence. Most shed leaves (>90%) were exported off the meadows by physical agents, such as waves and currents. The amount of belowground detritus stored within 10–15 cm from the sediment surface varied from ca. 70 to 7500 g DW m⁻² among the meadows examined, and they accumulated at rates ranging from ca. 65 to 650 g DW m⁻² per year. These values are large when compared to other communities of aquatic and terrestrial macrophytes. Our results show that P. oceanica meadows in the Spanish Mediterranean support high values of secondary production in other systems by exporting large amounts of leaf detritus as well as acting as substantial carbon sinks by accumulating large reservoirs of belowground detritus. Therefore, the increasing anthropogenic threats on P. oceanica could entail an important loss of secondary production and carbon storage in Mediterranean coastal ecosystems.     

Memorization of nuclear atmospheric tests by rhizomes and scales of the mediteranean seagrass Posidonia oceanica (Linnaeus) Delile

Determination of the age of rhizome sections of Posidonia oceanica (Linnaeus) Delile (Potamogetonaceae) by the examination of their dead leaf scales (lepidochronology), makes it possible to cut out the rhizomes in precisely dated 1- or 5-year sections. With increasing age of the section, the annual section length and the C:N ratio increase, while the dry weight per unit length and the ratio of dry weight:ash weight decrease. At a site where the only significant ¹³⁷Cs contamination can be ascribed to global nuclear fallout bombs, the activity of ¹³⁷Cs was measured i n 5-year age groups of rhizomes and scales separately. The maximum activity of ¹³⁷Cs in scales occurred in groups produced between 1960 and 1964, a period during which a peak of activity occurred in fallout. The distribution of ¹³⁷Cs activity in rhizome age groups indicates an apparent lag, perhaps owing to transport of material in the rhizome. In situ dead rhizome scales of P. oceanica thus constitute a memory, at least 30-years long, of an environmental chemical pulse. Posidonia oceanica could prove a valuable tool for marine-pollution surveys.     

Patch dynamics of the Mediterranean seagrass Posidonia oceanica: Implications for recolonisation process

Patch dynamics of the Mediterranean slow-growing seagrass Posidonia oceanica was studied in two shallow sites (3–10 m) of the Balearic Archipelago (Spain) through repeated censuses (1–2 year⁻¹). In the sheltered site of Es Port Bay (Cabrera Island), initial patch density (October 2001) was low: 0.05 patches m⁻², and the patch size (number of shoots) distribution was bimodal: most of the patches had less than 6 shoots or between 20 and 50 shoots. Mean patch recruitment in Es Port Bay (0.006 ± 0.002 patches m⁻² year⁻¹) exceeded mean patch loss (0.001 ± 0.001 patches m⁻² year⁻¹), yielding positive net patch recruitment (0.004 ± 0.003 patches m⁻² year⁻¹) and a slightly increased patch density 3 years later (July 2004, 0.06 patches m⁻²). In the exposed site of S’Estanyol, the initial patch density was higher (1.38 patches m⁻², August 2003), and patch size frequency decreased exponentially with size. Patch recruitment (0.26 patches m⁻² year⁻¹) and loss (0.24 patches m⁻² year⁻¹) were high, yielding a slightly increased patch density in the area 1 year later (October 2004, 1.40 patches m⁻²). Most recruited patches consisted of rooting vegetative fragments of 1–2 shoots. Seedling recruitment was observed in Summer 2004 at both sites. Episodic, seedling recruitment comprised 30% and 25% of total patch recruitment in Es Port Bay and S’Estanyol, respectively. Patch survival increased with patch size and no direct removal was observed among patches of 5 shoots or more. Most patches grew along the study, shifting patch distribution towards larger sizes. Within the size range studied (1–150 shoots), absolute shoot recruitment (shoots year⁻¹) increased linearly with patch size (R² = 0.64, p < 4 × 10⁻⁵, N = 125), while specific shoot recruitment was constant (about 0.25 ± 0.05 year⁻¹), although its variance was large for small patches. Given the slow growth rate and the high survival of patches with 5 or more shoots, even the low patch recruitment rates reported here could play a significant role in the colonisation process of P. oceanica.     

Shade-induced response and recovery of the seagrass Posidonia sinuosa

The effect of shading on the seagrass Posidonia sinuosa Cambridge et Kuo was investigated to identify mechanisms that prolong its survival during periods of low light and permit its subsequent recovery. We also tested whether the responses were consistent in plants growing at different depths. Shade treatments were low (LS; 70 – 100% of ambient Photosynthetic Photon Flux Density), medium (MS; 12 – 39%) and heavy (HS; 5 – 4%) at the shallow (3 – 4 m) site, whilst the deep (7 – 8 m) site had no HS treatment. HS at the shallow and MS at the deep site were below minimum light requirements (MLR) for the long-term survival of P. sinuosa. Physiological, morphological and growth attributes were repeatedly measured during 198 d of shade treatments and a subsequent 384 d recovery period at ambient PPFD. Shoot density declined by 82% within 105 d under HS treatment, though 6% of shoots remained after 198 d. We estimate that complete shoot loss in HS would have taken 2 years. Rhizome sugar concentrations declined to 32 – 52% of the controls at the end of the most severe shading treatments but after shoot loss, sugar concentrations declined more slowly or increased, suggesting a return to positive carbon balance. In the treatments below MLR, shading induced changes in physiological, morphological and growth characteristics, including reduced leaf length and width, reduced δ¹³C and photosynthetic adaptation to low light (increased α, reduced E_k and ETR_max), though not consistently. After removal of shading, photosynthetic characteristics became more typical of high light adaptation, possibly induced by greater light penetration through the thinned canopy, including reversal of the changes in α, E_k and ETR_max and induction of non-photochemical quenching. Carbohydrate concentrations increased to ambient concentrations within 115 d at ambient PPFD. Recovery of shoot density was slow, remaining significantly reduced in the MS and HS treatments after 384 d recovery. Shoot density at the end of shading is an important determinant of the rate seagrass meadows will recover and we estimated that the moderately and heavily shaded meadows would require 3.5 to 5 years to recover.     

Timing and success of reproductive stages in the seagrass Zostera noltii

The timing and success of sexual reproduction of the seagrass Zostera noltii was investigated at the Ria Formosa lagoon, Portugal. Thirty plants were tagged and monitored individually through time to determine in situ the development time of each maturation stage, from the emergence of the flowers to the production of seeds. The overall process of flowering and fruiting lasted 47 ± 4 days, during which formation and maturation of the fruits was the most time-consuming stage (27 ± 2 days). Spathe success, i.e. the percentage of spathes that produced seeds, was 22 ± 4% while spathe mortality was 34 ± 6%. A considerable percentage of spathes (37 ± 7%) was lost through leaf detachment, but some of these may still add to the reproductive success of the species by contributing to the species recruitment within the source meadow or elsewhere. Meadow seed production (MSP) of Z. noltii was estimated to be 312 ± 66 seeds m⁻², whereas the potential seed production of the studied meadow was 2623 seeds m⁻². Under laboratory conditions, 70% of Z. noltii seeds germinated within 26 days, but only 10% reached the seedling stage due to the high mortality of germlings. Fertility, defined as the probability of a seed to originate a new plant, was estimated to be 14 × 10⁻⁴, which is higher than what is expected for most seagrasses.     

Posidonia oceanica seedling root structure and development

The seedling root system of the seagrass Posidonia oceanica consists of a primary root and up to four adventitious roots. Under culture, germination and early growth began with the emergence of the primary root in the first week. Then the two adventitious root primordia originally present in the seed emerged at 3 and 5 weeks respectively, followed successively by further adventitious roots. Primary roots reached 17 mm at 4 weeks, but then their growth decreased markedly. In contrast the adventitious roots showed a pattern of continued elongation. Anatomical observations of both primary and adventitious roots revealed a multilayered hypodermis of thick-walled cells enclosing a wide cortex (99% of the root transverse area) and narrow stele. A well-distinguished endodermis was only observed in the primary roots, while differentiated xylem elements were found solely in the adventitious roots, but it is unclear to what degree differences between the two root types are due to different root maturity or to their role in water and nutrient uptake. Overall, the P. oceanica seedling root system is composed of multiple, rapidly formed roots which are strong yet flexible due to a large proportion of cortical tissue and further strengthened by a multilayered hypodermis, characteristics which could potentially facilitate initial anchorage and establishment.     

Effects of burial and erosion on the seagrass Zostera noltii

The effects of experimental burial and erosion on the seagrass Zostera noltii were assessed through in situ manipulation of the sediment level (− 2 cm, 0 cm, + 2 cm, + 4 cm, + 8 cm and + 16 cm). Shoot density, leaf and sheath length, internode length, C and N content and carbohydrates of leaves and rhizomes were examined 1, 2, 4 and 8 weeks after disturbance. Both burial and erosion resulted in the decrease of shoot density for all the sediment levels. The threshold for total shoot loss was between 4 cm and 8 cm of burial, particularly during the 2nd week. A laboratory experiment confirmed that shoots did not survive more than 2 weeks under complete burial. There was no evidence of induced flowering by burial or erosion. As well, no clear evidence was found of sediment level effects on leaf and sheath length. Longer rhizome internodes were observed as a response to both burial and erosion, suggesting a plant attempt to relocate the leaf-producing meristems closer to sediment surface or in search of new sediment avoiding the eroded area. The C content of leaves and rhizomes, as well as the non-structural carbohydrates (mainly the starch in rhizomes), decreased significantly along the experimental period, indicating the internal mobilization of carbon to meet the plant demands as a consequence of light deprivation. The significant decrease of N content in leaves, and its simultaneous increase in rhizomes, suggests the internal translocation of nitrogen from leaves to rhizomes. About 50% of the N lost by the leaves was recovered by the rhizomes. Our results indicated that Z. noltii has a high sensitivity to burial and erosion disturbance, which should be considered in the management of coastal activities.     

Recovery and succession in a multi-species tropical seagrass meadow following experimental disturbance: the role of sexual and asexual reproduction

Recolonisation and succession in a multi-species tropical seagrass meadow was examined by creating gaps (50×50 cm) in the meadow and manipulating the supply of sexual and asexual propagules. Measurements of leaf shoot density and estimates of above-ground biomass were conducted monthly to measure recovery of gaps between September 1995 and November 1997. Measurements of the seeds stored in the sediment (seed bank) and horizontal rhizome growth of colonising species were also conducted to determine their role in the recovery process.Asexual colonisation through horizontal rhizome growth from the surrounding meadow was the main mechanism for colonisation of gaps created in the meadow. The seed bank played no role in recolonisation of cleared plots. Total shoot density and above-ground biomass (all species pooled) of cleared plots recovered asexually to the level of the undisturbed controls in 10 and 7 months, respectively. There was some sexual recruitment into cleared plots where asexual colonisation was prevented but seagrass abundance (shoot density and biomass) did not reach the level of unmanipulated controls. Seagrass species did not appear to form seed banks despite some species being capable of producing long-lived seeds.The species composition of cleared plots remained different to the undisturbed controls throughout the 26-month experiment. Syringodium isoetifolium was a rapid asexual coloniser of disturbed plots and remained at higher abundances than in the control treatments for the duration of the study. S. isoetifolium had the fastest horizontal rhizome growth of species asexually colonising cleared plots (6.9 mm day⁻¹). Halophila ovalis was the most successful sexual coloniser but was displaced by asexually colonising species. H. ovalis was the only species observed to produce fruits during the study.Small disturbances in the meadow led to long-term (>2 years) changes in community composition. This study demonstrated that succession in tropical seagrass communities was not a deterministic process. Variations in recovery observed for different tropical seagrass communities highlighted the importance of understanding life history characteristics of species within individual communities to effectively predict their response to disturbance. A reproductive strategy involving clonal growth and production of long-lived, locally dispersed seeds is suggested which may provide an evolutionary advantage to plants growing in tropical environments subject to temporally unpredictable major disturbances such as cyclones.     

Evidence of the active role of ligno-cellulosic enzymes of marine fungi in degradation of Posidonia oceanica leaves

Evidence is given of release of six enzymes activities generally associated with plant material degradation, during in vitro experiments with Corollospora maritima acting upon Posidonia oceanica leaves. The results obtained are discussed in the light of the present knowledge on the role of marine fungi in the biological breakdown and conversion of leaf material in marine ecosystem.     

Posidonia oceanica in the Marmara Sea

The seagrass Posidonia oceanica is a stenohaline species endemic to the Mediterranean Sea, where it normally lives at a salinity of between 36.5 and 39.5 ppt. Surveys carried out at the North-eastern distribution limits revealed large beds in the Dardanelles Strait and isolated beds in the Marmara Sea, where the salinity ranges between 21.5 and 28 ppt. Microsatellite analysis of these low-salinity tolerant P. oceanica beds, show different signs of genetic isolation: excess of heterozygosity and a presence of fixed alleles. These particularities are rarely found in the whole distributional range of the species. Moreover, all the populations considered in the analysis have a very low genetic diversity in comparison with most of the meadows sampled throughout the Mediterranean Sea. Taking into consideration the genetic data, rhizome expansion rate and the actual extent of the isolated beds in the Marmara Sea and knowing the reproductive rate and dissemination characteristics of P. oceanica, we hypothesize that the isolated population of the Marmara Sea has been established since the Middle Holocene, before the catastrophic intrusion of brackish water into the Marmara Sea and the strong and persistent flow coming from the Black Sea.     

Estimation of the relative importance of sexual and vegetative reproduction in the clonal woodland herb Anemone nemorosa

Anemone nemorosa is a perennial rhizomatous plant of European woodlands. The “probability of clonal identity” method estimated the relative proportion of sexual to vegetative reproduction in this species to be 4.4% from allozyme genotype distributions. This result is congruent with investigations on the germination, short-term demography, population genetics, and breeding system of this species, and supports the hypothesis that even low levels of seedling recruitment can maintain considerable intrapopulational genetic diversity. Received: 2 March 1998 / Accepted: 24 July 1998     

Grazing as a mediator for maintenance of offspring diversity: Sexual and clonal recruitment in alpine grassland communities

To understand the effects of grazing on grassland plants sexual and clonal recruitment, we conducted a demographic field investigation of species recruitment along a grazing gradient in the Tibetan alpine grassland. Grazing intensity had significant effects on quantity and diversity of sexual and clonal recruitment. Sexual recruitment increased significantly, but clonal offspring production decreased significantly with increased grazing intensity. Grazing intensity had different, significant effects on offspring recruitment of the various functional groups in the community, grasses (GG), sedges (SG), legumes (LG) and forbs (FG). Higher grazing intensity reduced offspring recruitment of GG and SG; it increased offspring recruitment of LG and FG. Seedlings were significantly more abundant in lightly grazed, moderately grazed and heavily grazed meadows than in non-grazed grasslands. Offspring diversity from sexual recruitment was significantly higher than that from clonal recruitment in grazed than in non-grazed grasslands. Our studies indicate that moderate grazing had positive effects on seedling recruitment and offspring diversity, but heavy gazing may alter community succession by affecting recruitment patterns among the four plant functional groups.     

Reproductive phenology of the Mediterranean seagrasses Posidonia oceanica (L.) Delile, Cymodocea nodosa (Ucria) Aschers., and Zostera noltii Hornem.

Investigations on the sexual reproduction of Posidonia oceanica (L.) Delile, Cymodocea nodosa (Ucria) Aschers. and Zostera noltii Hornem. were conducted in situ from 1979 to 1988 in beds around the island of Ischia. Seed germination and phenological features of seedlings of P. oceanica and C. nodosa were observed in situ and in the laboratory. The reproductive cycles showed, for each species, different timing of start and duration of flowering, fruiting and germination. Posidonia oceanica flowers always appeared in September in shallow stands and in November in stands deeper than 15 m. This time delay seems to be related to the different maximum summer temperatures found at those depths. This species did not show dormancy. Cymodocea nodosa flowering was always recorded after the minimum winter temperature (April–May); germination of the seeds followed after 8–10 months of dormancy. Zostera noltii flowers were always recorded in July, with fruiting in August. The life cycles of these species seem to be related to the annual temperature range rather than to other environmental parameters. The different reproductive strategies adopted by each species are discussed.     

Reproductive cycle of Pitcairnia flammea Lindl. (Bromeliaceae/Pitcairnioideae) in an insular Atlantic rainforest area in southeastern Brazil

We studied for 6 years the reproductive cycle of the bromeliad Pitcairnia flammea in the Atlantic Rainforest of Ilha Grande, an island located in the south of Rio de Janeiro state, southeastern Brazil. We marked 150 adult individuals of the species, for which we monthly observed the occurrence of reproductive characteristics. The phenological attributes were related to environmental factors such as photoperiod, rainfall and temperature. Additionally, we estimated a mortality rate that was related to the rainfall in the area. P. flammea possesses a seasonal reproduction pattern, with a relatively fixed and well-defined time span during the winter dry period; flowering precedes fruiting by approx. 1 month. Due to the occurrence of P. flammea preferably on boulders of river channels and banks mortality of the plants is highest when floods from strong rainfall remove the plants mechanically from their growing places.     

The importance of grazing intensity and frequency for physiological responses of the tropical seagrass Thalassia hemprichii

Seagrass grazing is an intrinsic disturbance in primarily tropical and subtropical areas. While there is a general parabolic response in seagrass growth to grazing intensity, there is less knowledge on the role of grazing frequency, as well as potential interactions between grazing intensity and frequency. This study experimentally investigated physiological responses in Thalassia hemprichii to simulated (leaf cutting) grazing regimes with different intensities (25% vs. 75%) and frequencies (1 times vs. 3 times) over 35 days in Chwaka Bay (Zanzibar, Tanzania). The results showed that the two high-intensity treatments (75% removal) had 37–41% lower growth rate than the low-intensity/low-frequency treatment, and rhizome sugar and starch content were both affected in a similar way. A 36% lower starch content in the simulated low-intensity/high-frequency regime (25% × 3) compared to the one of low-intensity/low-frequency (25% × 1) also shows an interaction between grazing intensity and frequency. This suggests that high-intensity (and to some extent frequency) grazing regimes, in comparison to low-intensity regimes, could negatively affect T. hemprichii growth, energy reserves, and thereby the ability to deal with additional stress like light limitation or grazing.     

Specific growth rate as a determinant of the carbon isotope composition of the temperate seagrass Zostera marina

The seasonal variability of specific growth rate and the carbon stable isotope ratio (δ¹³C) of leaf blades (δ¹³C_leaf) of a temperate seagrass, Zostera marina (within 10 days old) were measured simultaneously, together with the δ¹³C of dissolved inorganic carbon (δ¹³C_DIC) at three sites in the semi-closed Akkeshi estuary system, northeastern Japan, in June, September, and November 2004. The δ¹³C_leaf ranged from −16.2 to −6.3‰ and decreased from summer to winter. The simultaneous measurement of the δ¹³C_leaf, growth rate, and morphological parameters (mean leaf length and width, mean number of leaves per shoot, and sheath length) of the seagrass and δ¹³C_DIC in the surrounding water allowed us to compare directly the δ¹³C_leaf and specific growth rate of seagrass. The difference in the δ¹³C of seagrass leaves relative to the source DIC (Δδ¹³C_{leaf − DIC}) was the least negative (−11 to −7‰) in June at all three sites and became more negative (−17 to −8‰) as the specific growth rate decreased. This positive correlation between Δδ¹³C_{leaf − DIC} and specific growth rate can be used to diagnose the growth of seagrasses. Δδ¹³C_{leaf − DIC} changed by −1.7 ± 0.2‰ when the leaf specific growth rate decreased by 1% d⁻¹.     

Vegetative multiplication and patch colonisation of Asarum europaeum subsp. europaeum L. (Aristolochiaceae) inferred by a combined morphological and molecular study

Asarum europaeum subsp. europaeum (Aristolochiaceae) is a rhizomatous herb forming distinct patches in calcareous broadleaved forests. Within natural stands, patches were mapped. In two regions, at least four patches were dug out, and connections between leaf modules through rhizomatous spacers were checked for signs of clonal reproduction (decay, breaking). Modules were sampled for amplified fragment length polymorphism (AFLP) fingerprinting to test whether they represent unique genets or are merigenets of a larger genet (split up by clonal reproduction), respectively.Morphologically, merigenet-relationships were only revealed in few cases with disrupted spacers between modules. With the obtained AFLP profiles for two primer combinations, the samples could be assigned to genets; clonal descendants of the same genet were readily identified. In one patch analysed in detail, 18 samples from 17 unconnected “plants” belonged to only two genets, which were morphologically divided into two and 15 merigenets, respectively. These two genets probably belonged to different maternal lineages and came into contact after lateral spread from the established clones. They showed divergent affinities to samples from adjacent patches (which all represented unique genets).The findings support the suitability of the combined morpho-ecological and molecular approach: compared to either method alone, it allows a more detailed analysis and interpretation of the fine-scale clonal structure, patch colonisation and especially of vegetative multiplication (with morpho-ecological studies to discern clonal growth and clonal reproduction and AFLP fingerprinting for genet and merigenet identification, respectively).     

Growth strategy of the stoloniferous rattan Calamus javensis in Mt. Halimun, Java

Calamus javensis is a widespread understorey rattan in Southeast Asia and is one of the dominant rattan species in Mt. Halimun National Park, West Java. This species can establish clumps either sexually with seeds or clonality with stolons. To analyze the role of sexual reproduction and clonality for clump establishment in C. javensis, we studied the vegetative and genetic structures of the population. Totals of 2,777 ramets comprising 1,321 clumps were found in three 0.16 ha plots. The number of ramets per clump showed reverse J-shaped distribution with a maximum of 15. Clustering and stoloniferous ramets accounted for 62 and 20% of all ramets, respectively. This indicates that C. javensis is actively engaged in clustering clonal growth. The longest stem was 14 m although 75% of all stems were shorter than 1 m long. The stolon ranged from 6 to 538 cm in length, which decreased the local density of ramets. Using randomly amplified polymorphic DNA (RAPD) analysis, we obtained 56 polymorphic bands for 118 clumps sampled from one of the plots. All samples showed different RAPD banding patterns, except one pair at a distance of 225 cm from each other. The genetic similarity among samples closer than 10 m was significantly higher than the expected value from a purely random distribution. These results suggest that most separate clumps are established by recruitment from seeds, and most seeds dispersed near mother plants. Hence, in C. javensis, clonality is more a growth strategy to increase the size of the genet than a dispersion-propagation strategy to expand the habitat.