Patch age alters seagrass response mechanisms to herbivory damage.

Natural disturbances can produce a mosaic of seagrass patches of different ages, which may affect the response to herbivory. These pressures can have consequences for plant performance. To assess how seagrass patch age affects the response to herbivory, we simulated the effect of herbivory by clipping leaves of Halodule wrightii in patches of 2, 4 and 6 years. All clipped plants showed ability to compensate herbivory by increasing leaf growth rate (on average 4.5-fold). The oldest patches showed resistance response by increasing phenolic compounds (1.2-fold). Contrastingly, the concentration of phenolics decreased in the youngest patches (0.26-fold), although they had a similar leaf carbon content to controls. These results suggest that younger plants facing herbivory pressure reallocate their phenolic compounds towards primary metabolism. Results confirm the H. wrightii tolerance to herbivory damage and provides evidence of age-dependent compensatory responses, which may have consequences for seagrass colonization and growth in perturbed habitats.

Resistance and recovery of benthic marine macrophyte communities to light reduction: Insights from carbon metabolism and dissolved organic carbon (DOC) fluxes, and implications for resilience.

A crucial factor in the long-term survival of benthic macrophyte communities under light-reduction stress is how they balance carbon metabolism during photosynthesis and respiration. In turn, the dissolved organic carbon (DOC) released by these communities, which can be highly light-dependent, stands as a source of carbon, fuelling marine communities and playing an important role in the ocean carbon sequestration. This is the first study to evaluate light-reduction stress and recovery in the seagrass Zostera noltei and the macroalga Caulerpa prolifera. Light reduction led to a significant decrease in the production of both communities from autotrophic to heterotrophic. Results indicated that most of the DOC released by vegetated coastal communities comes from photosynthetic activity, and that the net DOC fluxes can be greatly affected by shading events. Finally, both communities showed resilience underpinned by high recovery but low resistance capacity, with C. prolifera showing the highest resilience to unfavourable light conditions.

Recent trend reversal for declining European seagrass meadows.

Seagrass meadows, key ecosystems supporting fisheries, carbon sequestration and coastal protection, are globally threatened. In Europe, loss and recovery of seagrasses are reported, but the changes in extent and density at the continental scale remain unclear. Here we collate assessments of changes from 1869 to 2016 and show that 1/3 of European seagrass area was lost due to disease, deteriorated water quality, and coastal development, with losses peaking in the 1970s and 1980s. Since then, loss rates slowed down for most of the species and fast-growing species recovered in some locations, making the net rate of change in seagrass area experience a reversal in the 2000s, while density metrics improved or remained stable in most sites. Our results demonstrate that decline is not the generalised state among seagrasses nowadays in Europe, in contrast with global assessments, and that deceleration and reversal of declining trends is possible, expectingly bringing back the services they provide.

Global and local disturbances interact to modify seagrass palatability.

Global change, such as warming and ocean acidification, and local anthropogenic disturbances, such as eutrophication, can have profound impacts on marine organisms. However, we are far from being able to predict the outcome of multiple interacting disturbances on seagrass communities. Herbivores are key in determining plant community structure and the transfer of energy up the food web. Global and local disturbances may alter the ecological role of herbivory by modifying leaf palatability (i.e. leaf traits) and consequently, the feeding patterns of herbivores. This study evaluates the main and interactive effects of factors related to global change (i.e. elevated temperature, lower pH levels and associated ocean acidification) and local disturbance (i.e. eutrophication through ammonium enrichment) on a broad spectrum of leaf traits using the temperate seagrass Cymodocea nodosa, including structural, nutritional, biomechanical and chemical traits. The effect of these traits on the consumption rates of the generalist herbivore Paracentrotus lividus (purple sea urchin) is evaluated. The three disturbances of warming, low pH level and eutrophication, alone and in combination, increased the consumption rate of seagrass by modifying all leaf traits. Leaf nutritional quality, measured as nitrogen content, was positively correlated to consumption rate. In contrast, a negative correlation was found between feeding decisions by sea urchins and structural, biomechanical and chemical leaf traits. In addition, a notable accomplishment of this work is the identification of phenolic compounds not previously reported for C. nodosa. Our results suggest that global and local disturbances may trigger a major shift in the herbivory of seagrass communities, with important implications for the resilience of seagrass ecosystems.

Interaction between Ammonium Toxicity and Green Tide Development Over Seagrass Meadows: A Laboratory Study.

Eutrophication affects seagrasses negatively by increasing light attenuation through stimulation of biomass of fast-growing, bloom-forming algae and because high concentrations of ammonium in the water can be toxic to higher plants. We hypothesized nevertheless, that moderate amounts of nitrophilic macroalgae that coexists with seagrasses under eutrophic conditions, can alleviate the harmful effects of eutrophication on seagrasses by reducing ammonium concentrations in the seawater to non-toxic levels because such algae have a very large capacity to take up inorganic nutrients. We studied therefore how combinations of different ammonium concentrations (0, 25 and 50 µM) and different standing stocks of macroalgae (i.e. 0, 1 and 6 layers of Ulva sp.) affected survival, growth and net production of the seagrass Zostera noltei. In the absence of Ulva sp., increasing ammonium concentrations had a negative influence on the performance of Z. noltei. The presence of Ulva sp. without ammonium supply had a similar, but slightly smaller, negative effect on seagrass fitness due to light attenuation. When ammonium enrichment was combined with presence of Ulva sp., Ulva sp. ameliorated some of negative effects caused by high ammonium availability although Ulva sp. lowered the availability of light. Benthic microalgae, which increased in biomass during the experiment, seemed to play a similar role as Ulva sp.--they contributed to remove ammonium from the water, and thus, aided to keep the ammonium concentrations experienced by Z. noltei at relatively non-toxic levels. Our findings show that moderate amounts of drift macroalgae, eventually combined with increasing stocks of benthic microalgae, may aid seagrasses to alleviate toxic effects of ammonium under eutrophic conditions, which highlights the importance of high functional diversity for ecosystem resistance to anthropogenic disturbance.

Spatial variability and response to anthropogenic pressures of assemblages dominated by a habitat forming seaweed sensitive to pollution (northern coast of Alboran Sea).

The Cystoseira ericaefolia group is conformed by three species: C. tamariscifolia, C. mediterranea and C. amentacea. These species are among the most important habitat forming species of the upper sublittoral rocky shores of the Mediterranean Sea and adjacent Atlantic coast. This species group is sensitive to human pressures and therefore is currently suffering important losses. This study aimed to assess the influence of anthropogenic pressures, oceanographic conditions and local spatial variability in assemblages dominated by C. ericaefolia in the Alboran Sea. The results showed the absence of significant effects of anthropogenic pressures or its interactions with environmental conditions in the Cystoseira assemblages. This fact was attributed to the high spatial variability, which is most probably masking the impact of anthropogenic pressures. The results also showed that most of the variability occurred on at local levels. A relevant spatial variability was observed at regional level, suggesting a key role of oceanographic features in these assemblages.

Application of the CARLIT index along a biogeographical gradient in the Alboran Sea (European Coast).

An index, based on littoral communities assemblages (CARLIT), was applied to assess the ecological status of Northwestern Mediterranean coastal waters, following the requirements of the European Water Framework Directive. The biogeographical particularities of the Alboran Sea suggested a reassessment of this index, and that was the main objective of this work. Due to these biogeographical particularities, two regions were proposed in the studied region, with new reference conditions for each region. Subsequently, by means of a multivariate analysis, littoral community abundances and the CARLIT index were compared with factors related to geomorphology, biogeography and anthropogenic pressures. Overall, the biogeographical component determined the distribution of littoral communities. In contrast, the ecological status yielded by the index only was significantly related to anthropogenic pressures. The results pointed out that the reassessment of the CARLIT index was suitable to evaluate the ecological status of the Alboran Sea.

Multi-metric index based on the seagrass Zostera noltii (ZoNI) for ecological quality assessment of coastal and estuarine systems in SW Iberian Peninsula.

The aim of this study was to develop an ecological quality index based on the seagrass Zostera noltii (ZoNI) according to the WFD requirements. Eleven Z. noltii meadows of SW Iberian Peninsula under contrasting levels of anthropogenic disturbance were considered: 5 sites in Ria Formosa (Portugal), and 6 sites in Spain (Huelva and Cadiz). Environmental quality was assessed through nutrients of the water column and seagrass variables from different organization levels; those variables were analyzed using PCA (47% of explained variance on the first component) to calculate the ecological quality ratio (that was significantly correlated to the environmental variables, R(2)=0.51, p<0.01) and the ecological quality status of the sites. As a result, 4 sites were classified as good, 6 sites as moderate and 3 sites as poor ecological status. The developed index ZoNI showed to be suitable to assess the ecological status of estuarine and coastal systems in SW Iberian Peninsula reflecting their water quality.

GROWTH DYNAMICS OF ULVA ROTUNDATA (CHLOROPHYTA) IN A FISH FARM: IMPLICATIONS FOR BIOMITIGATION AT A LARGE SCALE(1).

Changes in biomass of several macroalgae [Ulva rotundata Bliding; Gracilariopsis longissima (S. G. Gmel.) Steentoft, L. M. Irvine et Farnham; Ulva intestinalis L.; and Cladophora sp.] and marine plants (Zostera noltii and Ruppia cirrhosa) growing naturally in earthen ponds of a fish farm (Acuinova, San Fernando, Southern Spain) were recorded during a year. The farm is mainly devoted to the culture of gilthered seabream (Sparus aurata). The most conspicuous algal species thriving in the ponds was U. rotundata, which reached densities up to 600 g dry mass · m(-2) and produced up to 20.45 g C · m(-2) · d(-1) . Dissolved nutrients (phosphate and ammonium), tissue nutrient content, and growth rates of this species were estimated during 2001 and 2002. Evidence of natural biomitigation by U. rotundata when water circulates throughout the fish farm is presented. Due to the fish cultivation, both phosphate and ammonium increased as water circulated from the preculture ponds to the postculture ponds. As a consequence, U. rotundata tissue nitrogen (N) and phosphorus (P) increased from algae growing in preculture ponds to algae growing in the outflow channel, so that mean C:N:P ratio varied from 773:57:1 in preculture ponds to 567:64:1 in the outflow channel. Phosphorus limited growth of U. rotundata during the spring. As growth rates increased as a function of tissue P, data were fitted to the Droop equation. From this equation, the estimated maximal growth rate was 0.295 ± 0.041 d(-1) , the subsistence quota was 0.05 ± 0.01% P of dry mass, and the critical quota was 0.215% P of dry mass. The results suggest that management of the fish farm based on a large-scale integrated mariculture system of fish and macroalgae may increase the total ecological and economic benefits, both for the farm and for the environment.

Evidence for a plasmalemma-based CO2 concentrating mechanism in Laminaria saccharina.

A kinetic analysis of the photosynthesis inhibition by buffers allowed quantification of some components of the carbon concentrating mechanism (CCM) of the brown macroalga Laminaria saccharina. The CCM was based on the presence of acid regions outside the plasma membrane that increased the CO(2) concentration available for photosynthesis by 10-20 times above that of the bulk medium at alkaline pH. Furthermore, the results suggested that the CCM is located mainly on the cell membrane and not in the chloroplast, as suggested for most macroalgae. The degree of dissipation of the acid regions by a buffer was related to the buffer anion concentration (B(-)), estimated from the titration of the buffer from bulk medium pH to a pH endpoint value close to the first pK (a) of the carbonic acid system. A kinetic model describing the relationship between inhibition of photosynthesis by a buffer and B(-) was developed suggesting that buffers act as competitive inhibitors with IC(50) (the concentration of the buffer anion which reduces the reaction velocity by half) of 5.0 mol m(-3). This model can be used to estimate the inhibitory effect of any buffer on the photosynthesis of L. saccharina. Nevertheless, some buffers tested showed a lower effect than that predicted from the hyperbolic model suggesting that their strength as inhibitors depended on: (1) the pK (a) in relation to the first pK (a) of the carbonic acid system and (2) its molecular weight (i.e. its mobility).

Fire, rain and the selection of seeder and resprouter life-histories in fire-recruiting, woody plants.

Several Cape species of the genus Erica are known to present seeder and resprouter phenotypes, and this variation seems to have a genetic basis. Therefore, this genus provides ideal model systems for using to elucidate the evolution of nonsprouting or seeder and resprouter life-histories in woody, fire-recruiting plants. A simple simulation model was developed to identify, under life-history optimality, the ecological conditions (viz. rainfall conditions and fire frequency) that conferred a selective advantage to the seeder phenotype over the resprouter in a given Cape Erica species. The model illustrated that the seeder life-history was able to invade and replace a resprouter population only under a mild mediterranean climate, with short, moderate summer droughts. This simulation approach will contribute to a better understanding of the biogeographical pattern of seeder and resprouter lineages of one of the paradigmatic fynbos woody taxa throughout the Cape floristic region.

Growth, carbon allocation and proteolytic activity in the seagrass Zostera noltii shaded by Ulva canopies.

The effects of light reduction [100%, 25%, 10% and 1% mean daily-integrated photon irradiance (I0)] by Ulva rigida C. Agardh canopies on carbon balance, sugar-related enzymes and proteolytic activities of the seagrass Zostera noltii Hornem. were investigated. Shaded plants showed negative net growth and starch was mobilized in both above- and below-ground tissues. Sucrose declined in below-ground parts under severe light deprivation (10% and 1% I0), but was accumulated in above-ground parts. Mobilization of the non-structural carbohydrates (sucrose and starch) was explained by changes in activities of sucrose synthase (SuSy, EC 2.4.1.13) and sucrose-phosphate synthase (SPS, EC 2.4.2.24). Under severe light reduction, the capacity of above-ground tissues for sucrose formation and export declined, indicated by the lowest SPS activity. In contrast, severe light reduction increased the 'sink strength' of below-ground tissues, demonstrated by the highest SuSy activities, and diminished the capacity for sucrose resynthesis from starch breakdown, as the lowest SPS activity was observed under low light. These results suggest a cessation of sucrose transport throughout the plant under extreme light limitation, the carbon supply being dependent on the starch breakdown in each tissue. The response of Z. noltii to gradual light reduction was co-ordinated at the whole-plant level, since an enhancement of proteolytic activities induced by carbon starvation in both above- and below-ground tissues was also recorded during prolonged light deprivation. Therefore, carbon mobilization was accompanied by enhanced protein turnover and changes in metabolic pathways.