Chlorophyll a concentration in plankton, biomass of different taxonomic phytoplankton groups, dominating algae species, and some parameters of coastal surface waters of the Black Sea

Based on data obtained at three stations in coastal waters of the Black Sea off Sevastopol in 2000 and 2001, we present seasonal dynamics of the carbon to chlorophyll a ratio in nano- and microphy-toplankton. This parameter varied approximately tenfold throughout the year. Its maximum values (442-500) were obtained in summer (July), when Pyrrophyta dominated in phytoplankton. Minimum values (36-56) were observed in winter (December),when diatomaceous species predominated. We derive a regression relating the carbon to chlorophyll a ratio to proportion of Pyrrophyta in total phytoplankton biomass, doing so separately for warm and cold seasons. Regression equations demonstrate coupled action of irradiance, temperature, and nutrient availability on the carbon to chlorophyll a ratio. For Pyrrophyta phytoplankton assemblage R**2 = 0.95, and for diatomaceous one R**2 = 0.87.

Impacts of food availability and pCO2 on planulation, juvenile survival, and calcification of the azooxanthellate scleractinian coral Balanophyllia elegans

Ocean acidification, the assimilation of atmospheric CO2 by the oceans that decreases the pH and CaCO3 saturation state (Omega) of seawater, is projected to have severe adverse consequences for calcifying organisms. While strong evidence suggests calcification by tropical reef-building corals containing algal symbionts (zooxanthellae) will decline over the next century, likely responses of azooxanthellate corals to ocean acidification are less well understood. Because azooxanthellate corals do not obtain photosynthetic energy from symbionts, they provide a system for studying the direct effects of acidification on energy available for calcification. The solitary azooxanthellate orange cup coral Balanophyllia elegans often lives in low-pH, upwelled waters along the California coast. In an 8-month factorial experiment, we measured the effects of three pCO2 treatments (410, 770, and 1220 µatm) and two feeding frequencies (3-day and 21-day intervals) on "planulation" (larval release) by adult B. elegans, and on the survival, skeletal growth, and calcification of newly settled juveniles. Planulation rates were affected by food level but not pCO2. Juvenile mortality was highest under high pCO2 (1220 µatm) and low food (21-day intervals). Feeding rate had a greater impact on calcification of B. elegans than pCO2. While net calcification was positive even at 1220 µatm (~3 times current atmospheric pCO2), overall calcification declined by ~25-45%, and skeletal density declined by ~35-45% as pCO2 increased from 410 to 1220 µatm. Aragonite crystal morphology changed at high pCO2, becoming significantly shorter but not wider at 1220 µatm. We conclude that food abundance is critical for azooxanthellate coral calcification, and that B. elegans may be partially protected from adverse consequences of ocean acidification in habitats with abundant heterotrophic food.

Nutrient availability affects the response of juvenile corals and the endosymbionts to ocean acidification

The interactive effects of nutrient availability and ocean acidification on coral calcification were investigated using post-settlement juvenile corals of Acropora digitifera cultured in nutrient-sufficient or nutrient-depleted seawater for 4 d and then exposed to seawater with different partial pressure of carbon dioxide () conditions (38.8 or 92.5 Pa) for 10 d. After the nutrient pretreatment, corals in the high nutrient condition (HN corals) had a significantly higher abundance of endosymbiotic algae than did those in the low nutrient condition (LN corals). The high abundance of endosymbionts in HN corals was reduced as a result of subsequent seawater acidification, and the chlorophyll a per algal cell increased. The photosynthetic oxygen production rate by endosymbionts was enhanced by the acidified seawater regardless of the nutrient treatment, indicating that the reduction in endosymbiont density in HN corals due to acidification was compensated for by the increase in chlorophyll a per cell. Though the photosynthetic rate increased in the acidified conditions for both LN and HN corals, the calcification rate significantly decreased for LN corals but not for HN corals. The acquisition of nutrients from seawater, rather than the increase in alkalinity caused by photosynthesis, might effectively alleviate the negative response of coral calcification to seawater acidification, suggesting that the response of corals and their endosymbionts to ocean acidification can be influenced by nutrient conditions.

Experiment: Food availability outweighs ocean acidification effects in juvenile Mytilus edulis

Ocean acidification is expected to decrease calcification rates of bivalves. Nevertheless in many coastal areas high pCO2 variability is encountered already today. Kiel Fjord (Western Baltic Sea) is a brackish (12-20 g kg-1) and CO2 enriched habitat, but the blue mussel Mytilus edulis dominates the benthic community. In a coupled field and laboratory study we examined the annual pCO2 variability in this habitat and the combined effects of elevated pCO2 and food availability on juvenile M. edulis growth and calcification. In the laboratory experiment, mussel growth and calcification were found to chiefly depend on food supply, with only minor impacts of pCO2 up to 3350 µatm. Kiel Fjord was characterized by strong seasonal pCO2 variability. During summer, maximal pCO2 values of 2500 µatm were observed at the surface and >3000 µatm at the bottom. However, the field growth experiment revealed seven times higher growth and calcification rates of M. edulis at a high pCO2 inner fjord field station (mean pCO2 ca. 1000 µatm) in comparison to a low pCO2 outer fjord station (ca. 600 µatm). In addition, mussels were able to outcompete the barnacle Amphibalanus improvisus at the high pCO2 site. High mussel productivity at the inner fjord site was enabled by higher particulate organic carbon concentrations. Kiel Fjord is highly impacted by eutrophication, which causes bottom water hypoxia and consequently high seawater pCO2. At the same time, elevated nutrient concentrations increase the energy availability for filter feeding organisms such as mussels. Thus M. edulis can dominate over a seemingly more acidification resistant species such as A. improvisus. We conclude that benthic stages of M. edulis tolerate high ambient pCO2 when food supply is abundant and that important habitat characteristics such as species interactions and energy availability need to be considered to predict species vulnerability to ocean acidification.

Light availability determines susceptibility of reef building corals to ocean acidification

Elevated seawater pCO2, and in turn ocean acidification (OA), is now widely acknowledged to reduce calcification and growth of reef building corals. As with other environmental factors (e.g., temperature and nutrients), light availability fundamentally regulates calcification and is predicted to change for future reef environments alongside elevated pCO2 via altered physical processes (e.g., sea level rise and turbidity); however, any potential role of light in regulating the OA-induced reduction of calcification is still unknown. We employed a multifactorial growth experiment to determine how light intensity and pCO2 together modify calcification for model coral species from two key genera, Acropora horrida and Porites cylindrica, occupying similar ecological niches but with different physiologies. We show that elevated pCO2 (OA)-induced losses of calcification in the light (G L) but not darkness (G D) were greatest under low-light growth conditions, in particular for A. horrida. High-light growth conditions therefore dampened the impact of OA upon G L but not G D. Gross photosynthesis (P G) responded in a reciprocal manner to G L suggesting OA-relieved pCO2 limitation of P G under high-light growth conditions to effectively enhance G L. A multivariate analysis of past OA experiments was used to evaluate whether our test species responses were more widely applicable across their respective genera. Indeed, the light intensity for growth was identified as a significant factor influencing the OA-induced decline of calcification for species of Acropora but not Porites. Whereas low-light conditions can provide a refuge for hard corals from thermal and light stress, our study suggests that lower light availability will potentially increase the susceptibility of key coral species to OA.

Individual dinoflagellate cysts species of surface sediment samples from the Mediterranean Sea

To determine the relationship between the spatial dinoflagellate cyst distribution and oceanic environmental conditions, 34 surface sediments from the Eastern and Western Mediterranean Sea have been investigated for their dinoflagellate cyst content. Multivariate ordination analyses identified sea-surface temperature, chlorophyll-a , nitrate concentration, salinity, and bottom oxygen concentration as the main factors affecting dinoflagellate cyst distribution in the region. Based on the relative abundance data, two associations can be distinguished that can be linked with major oceanographic settings. (1) An offshore eastern Mediterranean regime where surface sediments are characterized by oligotrophic, warm, saline surface water, and high oxygen bottom water concentrations (Impagidinium species, Nematosphaeropsis labyrinthus, Pyxidinopsis reticulata and Operculodinium israelianum). Based on the absolute abundance, temperature is positively related to the cyst accumulation of Operculodinium israelianum. Temperature does not form a causal factor influencing the accumulation rate of the other species in this association. Impagidinium species and Nematosphaeropsis labyrinthus show a positive relationship between cyst accumulation and nitrate availability in the upper waters. (2) Species of association 2 have highest relative abundances in the Western Mediterranean Sea, Strait of Sicily/NW Ionian Sea, and/or the distal ends of the Po/Nile/Rhône River plumes. At these stations, surface waters are characterized by (relative to the other regime) higher productivity associated with lower sea-surface temperature, salinity, and lower bottom water oxygen concentrations (Selenopemphix nephroides, Echinidinium spp., Selenopemphix quanta, Quinquecuspis concreta, Brigantedinium spp. and Lingulodinium machaerophorum). Based on both the absolute and relative abundances, Selenopemphix nephroides is suggested to be a suitable indicator to trace changes in the trophic state of the upper waters. The distribution of Lingulodinium machaerophorum is related to the presence of river-influenced surface waters, notably the Nile River. We suggest that this species might form a suitable marker to trace past variations in river discharge, notably from the Nile.

Differential response of two Mediterranean cold-water coral species to ocean acidification

Cold-water coral (CWC) reefs constitute one of the most complex deep-sea habitats harboring a vast diversity of associated species. Like other tropical or temperate framework builders, these systems are facing an uncertain future due to several threats, such as global warming and ocean acidification. In the case of Mediterranean CWC communities, the effect may be exacerbated due to the greater capacity of these waters to absorb atmospheric CO2 compared to the global ocean. Calcification in these organisms is an energy-demanding process, and it is expected that energy requirements will be greater as seawater pH and the availability of carbonate ions decrease. Therefore, studies assessing the effect of a pH decrease in skeletal growth, and metabolic balance are critical to fully understand the potential responses of these organisms under a changing scenario. In this context, the present work aims to investigate the medium- to long-term effect of a low pH scenario on calcification and the biochemical composition of two CWCs from the Mediterranean, Dendrophyllia cornigera and Desmophyllum dianthus. After 314 d of exposure to acidified conditions, a significant decrease of 70 % was observed in Desmophyllum dianthus skeletal growth rate, while Dendrophyllia cornigera showed no differences between treatments. Instead, only subtle differences between treatments were observed in the organic matter amount, lipid content, skeletal microdensity, or porosity in both species, although due to the high variability of the results, these differences were not statistically significant. Our results also confirmed a heterogeneous effect of low pH on the skeletal growth rate of the organisms depending on their initial weight, suggesting that those specimens with high calcification rates may be the most susceptible to the negative effects of acidification.

Availability of sequences of aquatic and terrestrial taxa in genetic databases (GenBank and BOLD - the Barcode of Life Database) in 2010, 2012 and 2016

Correct species identifications are of tremendous importance for invasion ecology, as mistakes could lead to misdirecting limited resources against harmless species or inaction against problematic ones. DNA barcoding is becoming a promising and reliable tool for species identifications, however the efficacy of such molecular taxonomy depends on gene region(s) that provide a unique sequence to differentiate among species and on availability of reference sequences in existing genetic databases. Here, we assembled a list of aquatic and terrestrial non-indigenous species (NIS) and checked two leading genetic databases for corresponding sequences of six genome regions used for DNA barcoding. The genetic databases were checked in 2010, 2012, and 2016. All four aquatic kingdoms (Animalia, Chromista, Plantae and Protozoa) were initially equally represented in the genetic databases, with 64, 65, 69, and 61% of NIS included, respectively. Sequences for terrestrial NIS were present at rates of 58 and 78% for Animalia and Plantae, respectively. Six years later, the number of sequences for aquatic NIS increased to 75, 75, 74, and 63% respectively, while those for terrestrial NIS increased to 74 and 88% respectively. Genetic databases are marginally better populated with sequences of terrestrial NIS of plants compared to aquatic NIS and terrestrial NIS of animals. The rate at which sequences are added to databases is not equal among taxa. Though some groups of NIS are not detectable at all based on available data - mostly aquatic ones - encouragingly, current availability of sequences of taxa with environmental and/or economic impact is relatively good and continues to increase with time.

Habitat traits and food availability determine the response of marine invertebrates to ocean acidification

Energy availability and local adaptation are major components in mediating the effects of ocean acidification (OA) on marine species. In a long-term study, we investigated the effects of food availability and elevated pCO2 (ca 400, 1000 and 3000 µatm) on growth of newly settled Amphibalanus (Balanus) improvisus to reproduction, and on their offspring. We also compared two different populations, which were presumed to differ in their sensitivity to pCO2 due to differing habitat conditions: Kiel Fjord, Germany (Western Baltic Sea) with naturally strong pCO2 fluctuations, and the Tjärnö Archipelago, Sweden (Skagerrak) with far lower fluctuations. Over 20 weeks, survival, growth, reproduction and shell strength of Kiel barnacles were all unaffected by elevated pCO2, regardless of food availability. Moulting frequency and shell corrosion increased with increasing pCO2 in adults. Larval development and juvenile growth of the F1 generation were tolerant to increased pCO2, irrespective of parental treatment. In contrast, elevated pCO2 had a strong negative impact on survival of Tjärnö barnacles. Specimens from this population were able to withstand moderate levels of elevated pCO2 over 5 weeks when food was plentiful but showed reduced growth under food limitation. Severe levels of elevated pCO2 negatively impacted growth of Tjärnö barnacles in both food treatments. We demonstrate a conspicuously higher tolerance to elevated pCO2 in Kiel barnacles than in Tjärnö barnacles. This tolerance was carried-over from adults to their offspring. Our findings indicate that populations from fluctuating pCO2 environments are more tolerant to elevated pCO2 than populations from more stable pCO2 habitats. We furthermore provide evidence that energy availability can mediate the ability of barnacles to withstand moderate CO2 stress. Considering the high tolerance of Kiel specimens and the possibility to adapt over many generations, near future OA alone does not seem to present a major threat for A. improvisus

Comparative responses of two dominant Antarctic phytoplankton taxa to interactions between ocean acidification, warming, irradiance, and iron availability

We investigated the responses of the ecologically dominant Antarctic phytoplankton species Phaeocystis antarctica (a prymnesiophyte) and Fragilariopsis cylindrus (a diatom) to a clustered matrix of three global change variables (CO2, mixed-layer depth, and temperature) under both iron (Fe)-replete and Fe-limited conditions based roughly on the Intergovernmental Panel on Climate Change (IPCC) A2 scenario: (1) Current conditions, 39 Pa (380 ppmv) CO2, 50 µmol photons/m**2/s light, and 2°C; (2) Year 2060, 61 Pa (600 ppmv) CO2, 100 µmol photons/m**2/s light, and 4°C; (3) Year 2100, 81 Pa (800 ppmv) CO2, 150 µmol photons/m**2/s light, and 6°C. The combined interactive effects of these global change variables and changing Fe availability on growth, primary production, and cell morphology are species specific. A competition experiment suggested that future conditions could lead to a shift away from P. antarctica and toward diatoms such as F. cylindrus. Along with decreases in diatom cell size and shifts from prymnesiophyte colonies to single cells under the future scenario, this could potentially lead to decreased carbon export to the deep ocean. Fe : C uptake ratios of both species increased under future conditions, suggesting phytoplankton of the Southern Ocean will increase their Fe requirements relative to carbon fixation. The interactive effects of Fe, light, CO2, and temperature on Antarctic phytoplankton need to be considered when predicting the future responses of biology and biogeochemistry in this region.

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.

Forest connectivity restoration through reforestation: an integrated methodology for prioritizing agricultural lands and selecting reforestation species

Forest connectivity restoration is a major goal in natural resource planning. Given the high amount of abandoned cultivated lands, setting efficient methods for the reforestation of agricultural lands offers a good opportunity to face this issue. However, reforestations must be carefully planned, which poses two main challenges. In first place, to determine those agricultural lands that, once reforested, would meet more effectively the planning goals. As a further step, in order to grant the success of the activity, it is fairly advisable to select those tree species that are more adapted to each particular environment. Here we intend to give response to both requirements by proposing a sequential and integrated methodology that has been implemented in two Spanish forest districts, which are formed by several landscape types that were previously defined and characterized. Using the software Conefor Sensinode, a powerful tool for quantifying habitat availability that is based on graph theory concepts, we determined the landscapes where forest planning should have connectivity as a major concern and, afterwards, we detected the agricultural patches that would contribute most to enhance connectivity if they were reforested. The subsequent reforestation species assessment was performed within these priority patches. Using penalized logistic regressions we fitted ecological niche models for the Spanish native tree species. The models were trained with species distribution data from the Spanish Forest Map and used climatic and lithological variables as predictors. Model predictions were used to build ordered lists of suitable species for each priority patch. The lists include dominant and non dominant tree species and allow adding biodiversity goals to the reforestation planning. The result of this combined methodology is a map of agricultural patches that would contribute most to uphold forest connectivity if they were reforested and a list of suitable tree species for each patch ordered by occurrence probability. Therefore the proposed methodology may be useful for suitable and efficient forest planning and landscape designing.

The relevance of seed size in modulating leaf physiology and early plant performance in two tree species

he size of seeds and the microsite of seed dispersal may affect the early establishment of seedlings through different physiological processes. Here, we examined the effects of seed size and light availability on seedling growth and survival, and whether such effects were mediated by water use efficiency. Acorns of Quercus petraea and the more drought-tolerant Quercus pyrenaica were sowed within and around a tree canopy gap in a sub-Mediterranean forest stand. We monitored seedling emergence and measured predawn leaf water potential (Ψpd), leaf nitrogen per unit area (Na), leaf mass per area, leaf carbon isotope composition (δ13C) and plant growth at the end of the first summer. Survival was measured on the next year. Path analysis revealed a consistent pattern in both species of higher δ13C as Ψpd decreased and higher δ13C as seedlings emerged later in the season, indicating an increase in 13C as the growing season is shorter and drier. There was a direct positive effect of seed size on δ13C in Q. petraea that was absent in Q. pyrenaica. Leaf δ13C had no effect on growth but the probability of surviving until the second year was higher for those seedlings of Q. pyrenaica that had lower δ13C on the first year. In conclusion, leaf δ13C is affected by seed size, seedling emergence time and the availability of light and water, however, leaf δ13C is irrelevant for first year growth, which is directly dependent on the amount of seed reserves.

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.

Selection of species and provenances for low-rainfall areas: physiological responses of Eucalyptus cloeziana and Eucalyptus argophloia to seasonal conditions in subtropical Queensland

Responses of stomatal conductance (g(s)) and net photosynthesis (A) to changes in soil water availability, photosynthetic photon flux density (Q), air temperature (1) and leaf-to-air vapour pressure deficit (D) were investigated in 4-year-old trees of a dry inland provenance of Eucalyptus argophloia Blakely, and two dry inland provenances (Coominglah and Hungry Hills) and a humid coastal provenance (Wolvi) of Eucalyptus cloeziana F. Muell. between April 2001 and April 2002 in southeast Queensland, Australia. There were minimal differences in A, g, and water relations variables among the coastal and inland provenances of E. cloeziana but large differences between E. argophloia and E. cloeziana. E. argophloia and to a lesser extent the Hungry Hills (inland) provenance of E. cloeziana maintained relatively higher pre-dawn water potential (psi(pd)) during the dry season suggesting possible access to water at depth. Simple phenomenological models of stomatal conductance as a function of Q, T and D explained 60% of variation in gs in E. cloeziana and more than 75% in E. argophloia, when seasonal effect was incorporated in the model. A Ball-Berry model for net photosynthesis explained between 70 and 80% of observed variation in A in both species. These results have implications in matching the dry and humid provenances of E. cloeziana and E. argophloia to suitable sites in subtropical environments. (C) 2004 Elsevier B.V. All rights reserved.