Poor success of seagrass Posidonia oceanica transplanting in a meadow disturbed by power line burial.

Local disturbances drive the decrease of the area covered by Posidonia oceanica in the Mediterranean. Mechanical impacts during the development of coastal infrastructures alter sea floor and the recolonization of benthic community will depend on the recovery of pre-disturbance environmental conditions and on the intrinsic characteristics of the local community that was disturbed. We transplanted 468 rhizome fragments and 450 seedlings of P. oceanica in a meadow disturbed by the trenching and deployment of a power line to evaluate the suitability of the disturbed sea floor for rehabilitating P. oceanica meadows. We quantify and compare the survivorship and vegetative development of the transplanted/planted (i.e. fragments/seedlings) material in the two types of the unconsolidated substrata left after infrastructure deployment works finished: sand and burlap bags filled with coarse gravel. The latter was used as a corrective measure for topographic restoration. Three experimental plots with sixteen transplanted fragments or twenty-five seedlings were placed at each substratum type at three different depths (i.e. 15, 20 and 25 m). Our results show that the transplanting of P. oceanica rhizome fragments in the disturbed substrata had low survival rates (0-31%) after 40-48 months. The survivorship of seedlings was lower than that of fragments. Our results highlight the importance of substratum for P. oceanica recovery after mechanical impact; disturbed, non-consolidated substrata will preclude P. oceanica rehabilitation through planting. Preservation of meadow substratum (i.e. dead matte) is a critical element that coastal infrastructure projects should consider to enable future recovery of P. oceanica meadows.

The role of competition and herbivory in biotic resistance against invaders: a synergistic effect.

Invasive species pose a major threat to global diversity, and once they are well established their eradication typically becomes unfeasible. However, certain natural mechanisms can increase the resistance of native communities to invaders and can be used to guide effective management policies. Both competition and herbivory have been identified as potential biotic resistance mechanisms that can limit plant invasiveness, but it is still under debate to what extent they might be effective against well-established invaders. Surprisingly, whereas biotic mechanisms are known to interact strongly, most studies to date have examined single biotic mechanisms separately, which likely influences our understanding of the strength and effectiveness of biotic resistance against invaders. Here we use long-term field data, benthic assemblage sampling, and exclusion experiments to assess the effect of native assemblage complexity and herbivory on the invasion dynamics of a successful invasive species, the alga Caulerpa cylindracea. A higher complexity of the native algal assemblage limited C. cylindracea invasion, probably through competition by canopy-forming and erect algae. Additionally, high herbivory pressure by the fish Sarpa salpa reduced C. cylindracea abundance by more than four times. However, long-term data of the invasion reflects that biotic resistance strength can vary across the invasion process and it is only where high assemblage complexity is concomitant with high herbivory pressure, that the most significant limitation is observed (synergistic effect). Overall, the findings reported in this study highlight that neglecting the interactions between biotic mechanisms during invasive processes and restricting the studied time scales may lead to underestimations of the true capacity of native assemblages to develop resistance to invaders.

Partitioning resilience of a marine foundation species into resistance and recovery trajectories.

The resilience of an ecological unit encompasses resistance during adverse conditions and the capacity to recover. We adopted a 'resistance-recovery' framework to experimentally partition the resilience of a foundation species (the seagrass Cymodocea nodosa). The shoot abundances of nine seagrass meadows were followed before, during and after simulated light reduction conditions. We determined the significance of ecological, environmental and genetic drivers on seagrass resistance (% of shoots retained during the light deprivation treatments) and recovery (duration from the end of the perturbed state back to initial conditions). To identify whether seagrass recovery was linearly related to prior resistance, we then established the connection between trajectories of resistance and recovery. Finally, we assessed whether recovery patterns were affected by biological drivers (production of sexual products-seeds-and asexual propagation) at the meadow-scale. Resistance to shading significantly increased with the genetic diversity of the meadow and seagrass recovery was conditioned by initial resistance during shading. A threshold in resistance (here, at a ca. 70% of shoot abundances retained during the light deprivation treatments) denoted a critical point that considerably delays seagrass recovery if overpassed. Seed densities, but not rhizome elongation rates, were higher in meadows that exhibited large resistance and quick recovery, which correlated positively with meadow genetic diversity. Our results highlight the critical role of resistance to a disturbance for persistence of a marine foundation species. Estimation of critical trade-offs between seagrass resistance and recovery is a promising field of research to better manage impacts on seagrass meadows.

A multidisciplinary approach to identify priority areas for the monitoring of a vulnerable family of fishes in Spanish Marine National Parks.

BACKGROUND: Syngnathid fishes (Actinopterygii, Syngnathidae) are flagship species strongly associated with seaweed and seagrass habitats. Seahorses and pipefishes are highly vulnerable to anthropogenic and environmental disturbances, but most species are currently Data Deficient according to the IUCN (2019), requiring more biological and ecological research. This study provides the first insights into syngnathid populations in the two marine Spanish National Parks (PNIA-Atlantic- and PNAC-Mediterranean). Fishes were collected periodically, marked, morphologically identified, analysed for size, weight, sex and sexual maturity, and sampled for stable isotope and genetic identification. Due the scarcity of previous information, habitat characteristics were also assessed in PNIA. RESULTS: Syngnathid diversity and abundance were low, with two species identified in PNIA (Hippocampus guttulatus and Syngnathus acus) and four in PNAC (S. abaster, S. acus, S. typhle and Nerophis maculatus). Syngnathids from both National Parks (NP) differed isotopically, with much lower δ15N in PNAC than in PNIA. The dominant species were S. abaster in PNAC and S. acus in PNIA. Syngnathids preferred less exposed sites in macroalgal assemblages in PNIA and Cymodocea meadows in PNAC. The occurrence of very large specimens, the absence of small-medium sizes and the isotopic comparison with a nearby population suggest that the population of Syngnathus acus (the dominant syngnathid in PNIA) mainly comprised breeders that migrate seasonally. Mitochondrial cytochrome b sequence variants were detected for H. guttulatus, S. acus, and S. abaster, and a novel 16S rDNA haplotype was obtained in N. maculatus. Our data suggest the presence of a cryptic divergent mitochondrial lineage of Syngnathus abaster species in PNAC. CONCLUSIONS: This is the first multidisciplinary approach to the study of syngnathids in Spanish marine NPs. Habitat preferences and population characteristics in both NPs differed. Further studies are needed to assess the occurrence of a species complex for S. abaster, discarding potential misidentifications of genus Syngnathus in PNAC, and evaluate migratory events in PNIA. We propose several preferential sites in both NPs for future monitoring of syngnathid populations and some recommendations for their conservation.

Seagrass Cymodocea nodosa across biogeographical regions and times: Differences in abundance, meadow structure and sexual reproduction.

Seagrasses are key habitat-forming species of coastal areas. While previous research has demonstrated considerable small-scale variation in seagrass abundance and structure, studies teasing apart local from large-scale variation are scarce. We determined how different biogeographic scenarios, under varying environmental and genetic variation, explained variation in the abundance and structure (morphology and biomass allocation), epiphytes and sexual reproduction intensity of the seagrass Cymodocea nodosa. Regional and local-scale variation, including their temporal variability, contributed to differentially explain variation in seagrass attributes. Structural, in particular morphological, attributes of the seagrass leaf canopy, most evidenced regional seasonal variation. Allocation to belowground tissues was, however, mainly driven by local-scale variation. High seed densities were observed in meadows of large genetic diversity, indicative of sexual success, which likely resulted from the different evolutionary histories undergone by the seagrass at each region. Our results highlight that phenotypic plasticity to local and regional environments need to be considered to better manage and preserve seagrass meadows.

Herbivory and resource availability shift plant defense and herbivore feeding choice in a seagrass system.

Numerous hypotheses have been posited to explain the observed variation in plant defense strategies against herbivory. Under resource-rich environments, plants are predicted to increase their tolerance (limiting resource model; LRM) and, while the resource availability hypothesis (RAH) predicts a decrease in constitutive resistance in plant species growing in resource-rich environments, at the intraspecific level, plants are predicted to follow an opposite pattern (intraspecific RAH). Furthermore, the effect of multiple factors in modulating plant defense strategies has been scarcely explored and is more difficult to predict. Our aim was to understand how plant defense traits respond to herbivory, resource availability and their interactions, and to assess the effects on plant palatability. To this end, we performed an in situ factorial experiment at two sites simulating three herbivory levels and two nutrient availability conditions with the seagrass Posidonia oceanica. Additionally, we performed a series of feeding experiments with its two main herbivores. While plants decreased their constitutive resistance under nutrient fertilization (contrary to intraspecific RAH but in accordance to the RAH), and did not increase allocation to tolerance (likely due to resource limitation, LRM), simulated herbivory induced resistance traits. However, we found no interactive effects of nutrient fertilization and herbivory simulation on plant defense. Both herbivores responded similarly to changes in plant palatability, strongly preferring nutrient-enriched plants and non-clipped plants. This work highlights the need to better understand the drivers of plant defense intraspecific variability in response to resources, particularly in habitat-forming species where changes in plant traits and abundance will cascade onto associated species.

Future warmer seas: increased stress and susceptibility to grazing in seedlings of a marine habitat-forming species.

Increases in seawater temperature are expected to have negative consequences for marine organisms. Beyond individual effects, species-specific differences in thermal tolerance are predicted to modify species interactions and increase the strength of top-down effects, particularly in plant-herbivore interactions. Shifts in trophic interactions will be especially important when affecting habitat-forming species such as seagrasses, as the consequences on their abundance will cascade throughout the food web. Seagrasses are a major component of coastal ecosystems offering important ecosystem services, but are threatened by multiple anthropogenic stressors, including warming. The mechanistic understanding of seagrass responses to warming at multiple scales of organization remains largely unexplored, especially in early-life stages such as seedlings. Yet, these early-life stages are critical for seagrass expansion processes and adaptation to climate change. In this study, we determined the effects of a 3 month experimental exposure to present and predicted mean summer SST of the Mediterranean Sea (25°C, 27°C, and 29°C) on the photophysiology, size, and ecology (i.e., plant-herbivore interactions) of seedlings of the seagrass Posidonia oceanica. Warming resulted in increased mortality, leaf necrosis, and respiration as well as lower carbohydrate reserves in the seed, the main storage organ in seedlings. Aboveground biomass and root growth were also limited with warming, which could hamper seedling establishment success. Furthermore, warming increased the susceptibility to consumption by grazers, likely due to lower leaf fiber content and thickness. Our results indicate that warming will negatively affect seagrass seedlings through multiple direct and indirect pathways: increased stress, reduced establishment potential, lower storage of carbohydrate reserves, and increased susceptibly to consumption. This work provides a significant step forward in understanding the major mechanisms that will drive the capacity of seagrass seedlings to adapt and survive to warming, highlighting the potential additive effects that herbivory will have on ultimately determining seedling success.

Seagrass (Posidonia oceanica) seedlings in a high-CO2 world: from physiology to herbivory.

Under future increased CO2 concentrations, seagrasses are predicted to perform better as a result of increased photosynthesis, but the effects in carbon balance and growth are unclear and remain unexplored for early life stages such as seedlings, which allow plant dispersal and provide the potential for adaptation under changing environmental conditions. Furthermore, the outcome of the concomitant biochemical changes in plant-herbivore interactions has been poorly studied, yet may have important implications in plant communities. In this study we determined the effects of experimental exposure to current and future predicted CO2 concentrations on the physiology, size and defense strategies against herbivory in the earliest life stage of the Mediterranean seagrass Posidonia oceanica. The photosynthetic performance of seedlings, assessed by fluorescence, improved under increased pCO2 conditions after 60 days, although these differences disappeared after 90 days. Furthermore, these plants exhibited bigger seeds and higher carbon storage in belowground tissues, having thus more resources to tolerate and recover from stressors. Of the several herbivory resistance traits measured, plants under high pCO2 conditions had a lower leaf N content but higher sucrose. These seedlings were preferred by herbivorous sea urchins in feeding trials, which could potentially counteract some of the positive effects observed.

Response of sulfate-reducing bacteria to an artificial oil-spill in a coastal marine sediment.

In situ mesocosm experiments using a calcareous sand flat from a coastal area of the island of Mallorca in the Mediterranean Sea were performed in order to study the response of sulfate-reducing bacteria (SRB) to controlled crude oil contamination, or heavy contamination with naphthalene. Changes in the microbial community caused by the contamination were monitored by a combination of comparative sequence analysis of 16S rRNA genes, fluorescence in situ hybridization, cultivation approaches and metabolic activity rates. Our results showed that crude oil and naphthalene negatively influenced the total microbial community as the natural increase in cell numbers due to the seasonal dynamics was attenuated. However, both contaminants enhanced the sulfate reduction rates, as well as the culturability of SRB. Our results suggested the presence of autochthonous deltaproteobacterial SRBs that were able to degrade crude oil or polycyclic aromatic hydrocarbons such as naphthalene in anaerobic sediment layers.

Antioxidant response of the seagrass Posidonia oceanica when epiphytized by the invasive macroalgae Lophocladia lallemandii.

The aim was to evaluate the antioxidant defences and the occurrence of oxidative damage in Posidonia oceanica under a stress situation due to the epiphytism of the invasive macroalgae Lophocladia lallemandii. P. oceanica leaves were collected in the absence (control station) and in the presence of the epiphytic algae L. lallemandii and the antioxidant enzyme activities, markers of oxidative damage, and hydrogen peroxide production were determined. Antioxidant enzyme--catalase, glutathione peroxidase and superoxide dismutase--activities were significantly higher in Posidonia epiphytized by L. lallemandii. Malondialdehyde, protein carbonyl derivates, and glutathione levels were also higher in L. lallemandii epiphytized P. oceanica leaves compared to control samples. The production of hydrogen peroxide was also significantly increased when Posidonia was epiphytized by L. lallemandii. The invasion of P. oceanica meadows by L. lallemandii appeared to induce oxidative stress in the seagrass as evidenced by increased levels of oxidative stress markers and antioxidant defences.

Biogeochemical conditions in sediments enriched by organic matter from net-pen fish farms in the Bolinao area, Philippines.

Sedimentation and sediment metabolism was measured at eight active milkfish fish pens and at one abandoned site in the Bolinao area, Philippines in order to examine the interactions between sediment and water in this shallow coastal zone. The rates of sedimentation were high in the area due to siltation, but the activities in the fish pens also contributed to enhanced sedimentation as indicated by the difference between the abandoned and active sites. The sediment metabolism appeared to decrease with increasing rates of sedimentation indicating that the microbial activity reached a saturation level in the fish pen sediments. Anaerobic processes dominated the organic matter decomposition, and sulfate reduction rates are among the highest measured in fish farm sediments. The rates decreased with increasing organic loading despite high concentrations of sulfate (>10 mM) at all sites. Presence of methane bubbles in the sediments suggests that sulfate reduction and methanogenesis were coexisting. The sediment metabolism was significantly reduced at the abandoned site indicating that the stimulation of microbial activities is due to active fish production. The anaerobic activity remained high at the abandoned site indicating that the sediment biogeochemical conditions remain affected long time after fish production has ceased.

Impacts of milkfish (Chanos chanos) aquaculture on carbon and nutrient fluxes in the Bolinao area, Philippines.

Sediment oxygen consumption, TCO2 production and nutrient fluxes across the sediment-water interface were measured in sediments within and along a transect from four fish pens with production of milkfish (Chanos chanos) in the Bolinao area, The Philippines. The four fish pens were each representing a specific period in the production cycling. There was a positive linear relationship between the rates of sedimentation inside the fish pens and the sediment oxygen consumption indicating that the benthic processes were controlled by the input of organic matter from fish production. The nutrient fluxes were generally higher inside the fish pens, and nitrate was taken up (1.7-5.8 mmol m(-2) d(-1)) whereas ammonium (1-22 mmol m(-2) d(-1)) and phosphate (0.2-4.7 mmol m(-2) d(-1)) were released from the sediments. The sediments were enriched in organic matter with up to a factor 4 compared to outside. A mass balance for one crop of milkfish was constructed based on production data and on measured fluxes of nutrients in the fish pens to assess the loss of carbon and nutrients to the environment. There was a loss to the surroundings of carbon and nitrogen of 51-68% of the total input, whereas phosphorus was buried in the sediments inside the fish pens which acted as net sinks of phosphorus. The results obtained suggest that fish pen culture as practiced in the Bolinao area, leads to even greater impacts on benthic carbon and nutrient cycling than those found in suspended cage cultures.