Leaf vs. epiphyte nitrogen uptake in a nutrient enriched Mediterranean seagrass (Posidonia oceanica) meadow

In situ nitrogen uptake by leaves and epiphytes was studied in a Mediterranean seagrass (Posidonia oceanica) meadow impacted from a fish farm and a pristine meadow, using ¹⁵NH₄ and ¹⁵NO₃ as tracers. In the impacted meadow both leaves and epiphytes yielded higher N concentrations and showed higher specific N uptake, suggesting a linkage between N uptake and its accumulation. Epiphytes took up N faster than leaves in relation to their corresponding biomass, but when assessed per unit area, N uptake was higher in leaves. Leaf N uptake was negatively correlated with epiphyte N uptake. With increasing epiphyte load on leaves, N leaf uptake decreased while N epiphyte uptake increased, indicating that epiphyte overgrowth hinders N uptake by P. oceanica leaves. Epiphyte contribution to total N uptake increased, while that of leaves decreased at the impacted meadow. However, 2-3 times less N was transferred daily from the water column to the benthic compartment, through seagrass and epiphyte uptake on total, at the impacted meadow. Therefore, it is probably still the loss of the key species - the seagrass - which plays the most important role in N cycling in seagrass ecosystems.     

Limited flexibility in resource use in a coral reef grazer foraging on seasonally changing algal communities

Feeding ecology of three life phases of the parrotfish Scarus ferrugineus was studied on a southern Red Sea fringing reef by comparing availability and consumption of benthic algae during the monsoon hot and cool seasons. Dominant biota covering dead carbonate substrates were in decreasing order of importance: turfs on endoliths, turfs on crustose corallines, and crustose corallines. On the reef crest and shallow fore reef, composition of the biota changed seasonally. Cover of turfs on endoliths and turfs on crustose corallines was higher during the hot season, while crustose corallines and macroalgae (only on reef crest) increased during the cool season. Biota in the deep fore reef did not show seasonal variation. All life phases used similar resources and showed selective feeding in all zones. Turfs on endoliths, followed by turfs on crustose corallines, was the primary feeding substrate. These two sources represented over 92% of bites during both seasons. Crustose corallines, macroalgae, and living corals were negligible components being strongly avoided at all zones and seasons. Resource use varied seasonally on the reef crest and shallow fore reef, while it remained unchanged on the deep fore reef. Turfs on endoliths were consistently preferred in both seasons but their contribution increased from 45% in the cool to 70% of bites in the hot season. Electivity for turfs on crustose corallines shifted from random feeding in the hot (27% of bites) to selection in the cool season (47% of bites). Feeding pattern changed diurnally with more bites taken from crustose corallines and turfs on crustose corallines during morning. During the rest of the day, bites from turfs on endoliths predominate. S. ferrugineus shows limited capacity to exploit seasonal increases in the biomass of foliose and canopy forming macroalgae, despite indications of energetic limitation during the cool season.     

Contributions of benthic and planktonic primary producers to nitrate and ammonium uptake fluxes in a nutrient-poor shallow coastal area (Corsica, NW Mediterranean)

By using the stable isotope ¹⁵N, we have measured in situ the uptake of nitrate and ammonium by the seagrass Posidonia oceanica, its leaf epiphyte community, the brown macroalgae Halopteris scoparia and the suspended particulate organic matter (SPOM). In Revellata Bay (Gulf of Calvi, Western Corsica), which is a very nutrient-poor region, the specific uptake rates (V) (μg N g N⁻¹ h⁻¹) of SPOM measured at ambient concentrations are 10-1000 higher than those of benthic primary producers. Macroalgae have intermediary V, between the seagrass leaf and leaf epiphytes. V are quite variable and the reasons for this variability remain unclear.Despite the difference of specific uptake rates found between benthic and pelagic primary producers, when integrating the uptake fluxes for a water column of 10 m depth, the contribution of benthic primary producers to N uptake fluxes (g N m⁻² h⁻¹) is significant, corresponding on average to 40% of total uptake flux. This results from the dominance in terms of N biomass of benthic primary producers in this shallow nutrient-poor area. When reported for the entire volume of the Revellata Bay, the contribution of benthic primary producers is reduced to 5-10% of total N uptake flux.Although this contribution could appear relatively low, it results in a significant direct transfer of inorganic nitrogen from the water column to the benthic compartment. By this transfer, the benthic plants act as a biological pump incorporating the pelagic N into the benthic compartment for a time longer than the characteristic time of phytoplankton dynamics (month-years vs. day-week).     

Effects of extreme seasonality on community structure and functional group dynamics of coral reef algae in the southern Red Sea (Eritrea)

Spatial and temporal variation in the biomass of four functional groups of coral reef algae (canopy algae, foliose algae, turf algae and crustose corallines) was investigated in the southern Red Sea. This region is characterised by extremely high summer temperatures (ca. 35°C). Strong seasonal shifts in the relative contribution of each group to the total macroalgal biomass were observed. On the reef flat, canopy and foliose algae dominated in winter, retaining low biomass in summer. On the fore reef, crustose corallines accounted for most of the macroalgal biomass throughout the year. Turf algae contributed least to the total biomass in all reef zones; biomass peaks shifted from midsummer on the inner reef flat to winter in the deeper zones. Biomass correlated negatively with seawater temperature in most groups, but the correlation was positive for turf algae on the shallow reef flat. We hypothesise both direct and indirect effects of the strong seasonality.

Occurrence of arsenic species in the seagrass <Emphasis Type="Italic">Posidonia oceanica</Emphasis> and in the marine algae <Emphasis Type="Italic">Lessonia nigrescens</Emphasis> and <Emphasis Type="Italic">Durvillaea antarctica</Emphasis>

The seagrass Posidonia oceanica from the Mediterranean coast and the brown algae Durvillaea antarctica and Lessonia nigrescens from the Chilean coast were analysed for total arsenic and for arsenic species. Arsenic species were extracted with water and measured by high-performance liquid chromatography–inductive coupled plasma mass spectrometry. Arsenite, arsenate, methylarsonate, dimethylarsinate, sulfonate sugar, sulphate sugar and phosphate sugar were analysed with an anion exchange column. Arsenobetaine, arsenocholine and glycerol sugar were separated with a cation exchange column. An extract of Fucus serratus was used to assign the arsenosugar peaks in the chromatograms from the sample extracts. In addition to the four common arsenosugars found in algae, inorganic arsenic was also measured in some samples, with the highest content found in L. nigrescens. The certified reference material (CRM) NIES no. 9 Sargassum fullvellum was analysed for quality assessment of total arsenic, and moreover, arsenosugars were also determined in this CRM.     

Some structural and functional aspects of the epiphytic component of four seagrass species (Cymodoceoideae) from Papua New Guinea

The epiphytic component of four monospecific seagrass beds from Papua New Guinea was studied structurally and functionally. The floristic composition and abundance of the epiphytes on leaves of four seagrass species (Cymodoceoideae) showed considerable variation, but on all four seagrass species, the same algae were among the five quantitatively most important epiphytes: encrusting coralline algae, Cyanophyta, Ceramium gracillimum (Harv.) Mazoyer, Polysiphonia savatierii Hariot and Audouinella spp. The temporal pattern of the epiphytic algae showed more or less the same features on the four seagrass species.Annual mean biomass of epiphytes and seagrass leaves ranged from 54 g ADW m^?2 in a community of Cymodocea rotundata Ehrenb. and Hempr. ex Aschers. to 169 g ADW m^?2 in a community of Syringodium isoetifolium (Aschers.) Dandy. The contribution of the epiphytic component to the total above-ground biomass ranged from 22 to 24%. Productivity of epiphytes was highest on leaves of Halodule uninervis (Forssk.) Aschers. (2.12 g ADW m^?2 sediment surface day^?1) and the epiphytic community contributed 35–44% of the total above-ground production of these four seagrass communities.     

The sedimentary facies of <Emphasis Type="Italic">Posidonia oceanica</Emphasis> seagrass meadows from the central Mediterranean Sea

Sedimentary facies of seven Posidonia oceanica meadows of western Mediterranean Sea were investigated. Five meadows are located in the Tyrrhenian coast, two are placed in the western coast of Sardinia and Corsica. These meadows develop on soft and hard substrates, often forming “mattes”, in areas characterized by different oceanography, morphology, and terrigenous inputs produced by coastal erosion and fluvial runoff. A total of five sedimentary facies have been recognized ranging from pure terrigenous to bioclastic: terrigenous sand to gravelly sand, bioclastic sands, skeletal gravelly sands, mixed siliciclastic–carbonate sands, well to moderately sorted skeletal siliciclastic sands. All of the sedimentary facies associated with P. oceanica are in the sand grain size. The gravelly fraction is generally subordinated and variable, whereas the muddy fraction is generally low. The very low frequencies of the muddy fraction can be attributed to re-suspension processes and to the lack of carbonate mud production. The rate of epiphytic carbonate production obtained by two of the investigated meadows averages 400 g m^?2 year^?1. This value is in the range of temperate Mediterranean as well as of tropical and subtropical seagrasses. The epiphytic carbonate production plus the calcareous biota living on seagrass substrate contributes to form mixed siliciclastic–carbonate sediments of the nearshore environment of the Mediterranean. Lastly, the carbonate production associated with seagrass was derived by biota belonging to the heterozoan assemblage, where aphotic organisms are dominant, together with oligophotic biota such as coralline algae and symbiont-bearing foraminifera. Consequently, in the well-illuminated seagrass settings, the prevalent skeletal assemblages is represented by the heterozoan association while the components of the photozoan assemblages are absent or subordinate. This a key point for the paleoenvironmental reconstruction of the photic zone in the fossil record. Because the skeletal components of many seagrass dwellers greatly contribute to the carbonate sediment production of photic shallow-water environments, the seagrass meadows became substantial places of carbonate production and C (organic and inorganic) sequestration during the Cenozoic.     

Physiological responses of intertidal marine brown algae to nitrogen deprivation and resupply of nitrate and ammonium

Intertidal macroalgae Fucus and Laminaria experience seasonally fluctuating inorganic N supply. This study examined the effects of long‐term N deprivation, recovery following N resupply, and effects of elevated ammonium and nitrate exposure on N acquisition in intertidal algae using manipulations of N supply in tank culture. Over 15 weeks of N deprivation, internal N and nitrate reductase activity (NRA) declined, but maximum quantum yield of PSII was unaffected in Fucus serratus and Fucus vesiculosus. Low NRA was maintained despite no external nitrate availability and depletion of internal pools, suggesting a constitutive NRA, insensitive to N supply. Nitrate resupplied to N‐starved thalli was rapidly taken up and internal nitrate pools and NRA increased. Exposure to elevated (50 μM) nitrate over 4 days stimulated nitrate uptake and NRA in Laminaria digitata and F. serratus. Exposure to elevated ammonium suppressed NRA in L. digitata but not in F. serratus. This novel insensitivity of NRA to ammonium in Fucus contrasts with regulation of NRA in other algae and higher plants. Ammonium suppression of NRA in L. digitata was not via inhibition of nitrate uptake and was independent of nitrate availability. L. digitata showed a higher capacity for internal nitrate storage when exposed to elevated ambient nitrate, but NRA was lower than in Fucus. All species maintained nitrate assimilation capacity in excess of nitrate uptake capacity. N uptake and storage strategies of these intertidal macroalgae are adaptive to life in fluctuating N supply, and distinct regulation of N metabolism in Fucus vs Laminaria may relate to position in the intertidal zone.     

Influence of ammonium and nitrate supply on growth, dry matter partitioning, N uptake and photosynthetic capacity of Pinus radiata seedlings

Growth and physiological responses of Pinus radiata D. Don seedlings to a combination of N supply regimes (low N = 1.78 mol m⁻³, high N = 7.14 mol m⁻³) and ammonium:nitrate ratios (80:20, 50:50 and 20:80; molar basis) were assessed in a hydroponic experiment run over the course of 105 days. Highly significant (P < 0.001) increases in seedling diameter, height, leaf area and dry mass occurred at lower ammonium:nitrate ratios and were two to fourfold greater than the non-significant (for diameter) to marginally significant (P < 0.05 for other dimensions) increases in these dimensions that occurred with greater N supply. Increases in N supply resulted in a highly significant (P < 0.001) reduction in biomass partitioning to roots and highly significant (P < 0.001) increases in allocation to foliage. The ammonium:nitrate ratio was not found to significantly change biomass partitioning to either foliage, stems or roots. Ammonium and nitrate uptake was significantly influenced by N supply and N form and conformed to ammonium and nitrate concentrations in nutrient solution. Uptake rates of ammonium were twice those of nitrate at comparable concentrations suggesting that P. radiata is in the lower end of the ratio of uptake of ammonium to nitrate reported for conifers (range from 2 to 20 mol mol⁻¹). Despite this, plants growing in high ammonium:nitrate ratios were smaller, exhibited luxurious N consumption and lower N use efficiency. Differences in productivity among treatments were partially explained by greater rates of light-saturated photosynthesis associated with nitrate nutrition.     

NITROGENOUS NUTRITION OF ALEXANDRIUM CATENELLA (DINOPHYCEAE) IN CULTURES AND IN THAU LAGOON,SOUTHERN FRANCE1

Alexandrium catenella (Whedon et Kofoid) Balech was isolated from Thau lagoon (northern Mediterranean) and its growth and uptake characteristics measured for nitrate, ammonium, and urea. Although affinity constants did not indicate a preference for ammonium over nitrate, there was a strong inhibition of nitrate uptake by ammonium when both nitrogen (N) sources were present. Nitrogen budgets during growth in cultures revealed major imbalances between decreases in dissolved N and increases in particulate N, indicating excretion of dissolved organic N during the early part of the growth phase and uptake during the later part. A quasi‐unialgal bloom in November 2001 (4×10⁶ cells·L^?1) allowed measurements of uptake of nitrate, nitrite, ammonium, and urea; net and gross growth rate of A. catenella; and grazing rates on this organism. The affinity constants indicate that it is not a strong competitor for the N nutrients tested when these are in low concentrations (<10 μgat N·L^?1), compared with other members of the phytoplankton community. Indirect evidence from cultures indicate that dissolved organic N compounds could be important in triggering those blooms. Finally, the strongly unbalanced growth observed in the field indicates that A. catenella exhibits a storage rather than a growth response to a nutrient pulse and is adapted to low frequency events such as the passage of frontal disturbances. The disappearance of A. catenella was due to grazing that balanced growth at the peak of the bloom.     

Effects of Posidonia Oceanica Beach-Cast on Germination,Growth and Nutrient Uptake of Coastal Dune Plants

Seagrass meadows play an important role in marine ecosystems. A part of seagrass production is also exported to adjacent coastal terrestrial systems, possibly influencing their functioning. In this work we experimentally analyzed the effect of Posidonia oceanica beach-cast on plant germination, growth, and nutrient uptake of two plant species (Cakile maritima and Elymus farctus) that grow on upper beaches and fore dunes along the Mediterranean coasts. We compared plants growing in simple sand (control) with those growing in a substrate enriched with P. oceanica wrack (treatment) in laboratory. P. oceanica wrack doubled the N substrate pool and kept the substrate humid. Plants growing in the treated substrate grew faster, were twice as large as those growing in the control substrate, while tissues were enriched in N and P (Cakile by the 1.3 fold in N and 2.5 fold in P; Elymus by 1.5 fold in N and 2 fold in P). Our results suggest a positive effect of seagrass litter for the enhancing of dune species, highlighting its role for the conservation of coastal dune ecosystems.     

The impact of nitrogen deposition on photobiont‐mycobiont balance of epiphytic lichens in subtropical forests of central China

Excessive nitrogen (N) deposition can impact lichen diversity in forest ecosystems, and this is a particular situation in China. Here, we examined the N uptake, assimilation, and the impact of excessive N deposition on the symbiotic balance of dominant epiphytic lichens in the subtropical forests in the Mts. Shennongjia of central China. The results show that lichen species took up, assimilated and utilized more ammonium than nitrate in a species‐specific way, following the increase of N availability. The photobiont of the lichens decreased with the increase of N concentration following an initial increase, while the mycobiont response to the N addition was not apparent. Considerable variation in response to excessive N deposition exists among the lichen species. Usnea longissima could regulate its N uptake, resulting in a stable photobiont‐mycobiont ratio among N treatments. In contrast, the photobiont‐mycobiont ratio of other four lichens increased initially but decreased when N concentration exceeded a certain level, and N stress may have broken the balance between photobiont and mycobiont of these lichens. Our results suggest that most epiphytic lichens in subtropical forest of central China could uptake and assimilate more ammonium than nitrate and that the balance between photobiont and mycobiont of many epiphytic lichens might change with the increasing N deposition load, which could impact the lichen diversity of this forest ecosystem.     

Molecular Characterization of a Variable Tandem Repeat Sequence Determined during RAPD Analysis among Posidonia oceanica Insular and Coastline Populations

The seagrass Posidonia oceanica plays a multifunctional role in the coastal area as an important and productive component of ecosystems in the Mediterranean Sea. We detected by RAPD analysis with two arbitrary primers genetic differences in P. oceanica collected from several sites in the Southern Mediterranean. By AMOVA analysis we observed a level of about 20% genetic difference among individuals within a population and 80% among populations. A common band of 200 bp was found in all the amplified samples. Cloning and sequencing analysis of this band revealed the presence of a simple tandem repeat sequence (minisatellite) that we called PoTR (Posidonia oceanica tandem repeat). Finally, the ability of PoTR to detect genetic variability inP. oceanica genome was demonstrated by the presence of amplification products of different lengths utilizing primers internal to this sequence.     

Seasonal shifts in seagrass bed primary producers in a cold-temperate estuary: Dynamics of eelgrass Zostera marina and associated epiphytic algae

The seasonal dynamics of seagrass and epiphytic algal primary production were measured in an eelgrass (Zostera marina) bed in the Akkeshi-ko estuary, Hokkaido, Japan (43°02′N, 144°52′E). During spring and early summer, eelgrass biomass increased, with a high production (maximum: 2.89 g C m⁻² day⁻¹), but the production and biomass of epiphytic algae remained low. In contrast, epiphytic algae bloomed in August, with a high production (5.21 g C m⁻² day⁻¹), but eelgrass production ceased and its biomass subsequently decreased. Therefore, the major primary producers in this eelgrass bed switched seasonally from eelgrass in spring and early summer to epiphytic algae in late summer and autumn. Epiphytic algae maintained similar productivity because of the change of photosynthetic kinetics and the dominant epiphytic diatom changed from highly adhesive species to less adhesive or filamentous small species during the bloom. This suggests that the change of epiphyte density and biomass was due to change of its loss rate, possibly due to herbivorous grazing rate. Moreover, competition between epiphytic algae and eelgrass for nutrients and light may also affect the dramatic seasonal changes in the major primary producers.     

Nutritional interaction in an alga-barnacle association

Nitrogen is the major growth-limiting nutrient for marine algae. One potential source of nitrogen for marine algae is ammonium released by invertebrates. Many mid-intertidal reefs in northeastern New Zealand are dominated by a close association between the honeycomb barnacle Chamaesipho columna and an encusting brown alga Pseudolithoderma sp. Growth of Pseudolithoderma was enhanced in the presence of live C. columna, which released ammonium at a greater rate than the maximum rate of ammonium uptake by Pseudolithoderma. Algal tissue on barnacle tests had a lower C:N ratio than tissue located more than 2 cm from the nearest barnacle, suggesting the barnacle is an important source of nitrogen for the alga. The role of nutrient exchange in determining ecological patterns of species in marine communities is discussed.     

Invasion of alien macroalgae in different Mediterranean habitats

The study evaluated different macroalgal invasions in the main Mediterranean coastal habitats on hard bottom. Biodiversity, species composition and structure of macroalgal assemblages were compared among non-invaded areas and areas invaded by the Chlorophyta Caulerpa racemosa var. cylindracea and by the turf-forming Rhodophyta Womersleyella setacea in three different habitats: shallow rocky bottom, deep rocky bottom and dead matte of the seagrass Posidonia oceanica. Results showed that alien macroalgae constituted a relevant component of benthic assemblages in invaded areas of the Mediterranean Sea. Assemblages invaded by Womersleyella setacea and Caulerpa racemosa showed lower values of diversity and large differences in the structure and species composition related to non-nvaded assemblages. The species that mostly suffered from invasion were erect species reproducing sexually; moreover, the dominance of W. setacea led to low abundance of native filamentous algae, while C. racemosa colonization seemed particularly threatening for encrusting algae. All the studied habitats appeared highly invasible by alien macroalgae, even if W. setacea appeared more invasive in deeper habitats, while colonization of C. racemosa seemed more serious in shallower habitats; the dead matte of P. oceanica represented a suitable substrate for the spread of both species. Differences among assemblages in different habitats were reduced in invaded areas.     

Grazing effects of a gammaridean Amphipoda,Ampithoe sp., on the seagrass,Syringodium isoetifolium,and epiphytes in a tropical seagrass bed of Fiji

In the pure stand of tropical seagrass,Syringodium isoetifolium, in a small oceanic island, Fiji, grazing effects of the seagrass-associated gammarid,Ampithoe sp., on seagrass and epiphytes were assessed in October 1989, November 1991, November 1992. Density of the gammarid was estimated with two methods, mesh bag method and tuft method. During the three years surveyed the density of the gammarid increased remarkably from 1989 to 1991, with heavy epiphytism. Gut contents of the gammarid were examined. Grazing rates on seagrass leaf with and without epiphytic blue-green algae were measured in a bottle experiment. Litter bag experiments were conducted using different mesh sizes each containing seagrass only and seagrass and gammarids. The seagrass leaf biomass in the litter bag reduced abruptly in both bags. After one week, 78–86% of seagrass biomass disappeared from the bags. Enhancement of decomposition of seagrass leaf by the gammarid grazing was observed. Oxygen consumption and ammonium excretion rates were measured simultaneously in bottle experiments. Carbon budget in the seagrass bed was estimated as follows: 0.9 gC m⁻² day⁻¹ in seagrass growth, gammarid grazing was about a half of it and further assimilated a half of it, about 0.1 gC m⁻² day⁻¹, and more than half of it become CO₂ by respiration. Grazing effects on epiphyte and seagrass growth and production were discussed through the carbon budget and indirect interactions between seagrass, epiphytes and associated gammarids to explain the temporal change of seagrass and epiphyte dynamics.     

Effects of an epiphytic alga on abundances of fish and decapods associated with the seagrass Zostera capricorni

During a field experiment to test effects of changes to leaf height and density of Zostera capricorni on associated fish and decapods, more than one-third of the substratum at some sites was covered by the epiphytic brown alga Giffordia mitchelliae. This paper reports apparent effects of the alga on the 13 most abundant species of fish and decapods, and interactions between effects of the alga and those due to height and density of the seagrass leaves. Abundances of three species were significantly lower, and those of two species were significantly greater where the alga occurred. There were interactions between effects of the alga and those of leaf height and/or density for another four species. In all four cases, when the alga had an effect, it was to decrease abundance. The effects of Giffordia were generally opposite to those reported for drift algae in seagrass habitats elsewhere. The lower abundances of seven species where algal cover was great are best explained by an existing model for seagrass habitats that predicts reductions in abundance of individuals when the ratio of plant material to substratum is very high.