Diversity and Distribution of the Bryozoa along Underwater Cliffs in the Adriatic Sea with Special Reference to Thermal Regime

Abstract.  The diversity and distribution of bryozoans have been surveyed at two localities, Cape Struga on Lastovo Island (South Adriatic) and Cape Šilo on Prvić Island near Senj (North Adriatic), with similar characteristics of topography and exposure but markedly different sea temperature conditions. Continuous measurements over one year have shown that temperature conditions differ in two aspects. First, the annual range of temperature is much larger in the North. Second, in the South during summer, marine life within the thermocline layer is exposed to large high-frequency temperature oscillations, partly due to an internal diurnal tide, a phenomenon recorded for the first time in the Adriatic. Altogether, 78 species have been recorded, 57 in Lastovo and 50 in Prvić, and 29 in both localities. The comparison of mean zooid lengths of 14 bryozoan species occurring at both localities indicates no consistent difference in zooid size between northern and southern sites. Seventeen bryozoan species have been recorded in the Eastern Adriatic Sea for the first time.     

Life history of Cornularia cornucopiae (Anthozoa: Octocorallia) on the Conero Promontory (North Adriatic Sea)

The life cycle of the stoloniferan Cornularia cornucopiae (Pallas, 1766) (Anthozoa: Octocorallia) was studied from March 2009 to October 2010 on the rocky cliff of the Conero Promontory (North Adriatic Sea, 43°34.865′ N, 13°34.320′ E). In this area the species showed unusual high densities never recorded in other sites of the Mediterranean Sea. The density trend of the species showed a marked seasonal cycle, with a winter minimum of about 1000 polyps m^?2 and a summer maximum of about 30,000 polyps m^?2. In accordance with other Mediterranean literature data, polyps were fertile during spring–summer, from March to August, but the number of eggs per polyp continuously decreased during this span of time. Variations of polyp density were strongly correlated to water temperature, which can be considered the main environmental factor triggering this seasonal behaviour. The possibility, for C. cornucopiae, to face adverse winter conditions is probably related to the presence of a characteristic perisarcal envelope covering the stolon and the calyx of each polyp, which isolates the living tissues from the exterior. During winter, polyps degenerate but the stolons remain dormant inside their envelopes. The perisarc covering represents a morphological convergence of C. cornucopiae with benthic hydrozoans. As the latter, the studied stoloniferans are able to live in habitats characterized by periodic favourable conditions thanks to a seasonal life strategy. A similar trend is shared also by other important components (cnidarians and some sponges) of the filter‐feeding community of the North Adriatic Sea. Differently to the Western Mediterranean basin, this area is characterized by high food availability all year around, so benthic organisms are strongly constrained by the very low winter temperatures.     

Effects of eelgrass (Zostera marina) canopy removal and sediment addition on sediment characteristics and benthic communities in the Northern Baltic Sea

The eelgrass Zostera marina is a key structural and functional species across the European coastline. The separate and interactive effects of eelgrass canopy removal and sediment addition on the sediment characteristics and the structure of benthic communities were studied in a factorial field experiment in the Northern Baltic Sea in July–August 2006. The removal of eelgrass canopy temporarily increased the sediment oxygen consumption, reduced the content of fine particles (<100 μm) and organic matter in the sediment, and increased the share of sand fraction (250–500 μm). Sediment addition increased the content of fine particles (<100 μm) and reduced the share of sand fraction (250–1000 μm). The effects were strongest in the presence of eelgrass canopy. Benthic invertebrates and macroalgae were affected by eelgrass canopy removal but not by sediment addition. The removal of eelgrass canopy significantly decreased benthic species richness and invertebrate and macroalgal densities. To conclude, our experiment demonstrates that Z. marina defines the patterns of benthic macroalgae and invertebrates but has moderate effects on sediment structure and metabolism in the Northern Baltic Sea.     

Population structure and conservation status of the red gorgonian Paramuricea clavata (Risso, 1826) in the Eastern Adriatic Sea

Gorgonians are important structuring species of the Mediterranean hard‐bottom communities that are threatened by disturbances such as increasing seawater temperature, mucilaginous events and destructive fishing, among others. In this study we assessed for the first time the population structure and conservation status of one of the most common gorgonians in the Eastern Adriatic Sea, the red gorgonian Paramuricea clavata. During late spring 2009, nine populations dwelling between 30 and 50 m depth were examined by SCUBA diving along 200 km of the Croatian coastline. The density ranged between 7 and 20 colonies·m^?2. The mean and maximum colony heights were 31.2 ± 22.7 cm (±SD) and 138 cm, respectively. Two main patterns of P. clavata size frequency distributions were observed: the first one with a higher proportion of juveniles (~30%) observed mostly in the northernmost populations, and the second one with a higher proportion of larger colonies (>25% of colonies >40 cm in height). Regarding the disturbance impact level, the proportion of healthy colonies (with <10% of injured surface) was high in almost all of the studied populations (>60%) and the mean extent of injury (i.e. denuded axis or epibiosis) was 9.7 ± 4% (±SD), indicating low impacts. Contrasting population size structures with high recruitment in mature populations provides new insights into the demographic structure of the Mediterranean gorgonian forests dwelling in their upper bathymetric range (<50 m depth). Furthermore, these size structures and the low impact levels suggest a current favorable conservation status of the studied populations in the Eastern Adriatic Sea and provide a baseline for their monitoring in the future.     

Trends in oxygen content 1911–1984 and occurrence of benthic mortality in the northern Adriatic Sea

Trends of dissolved oxygen content of the surface and bottom layers of the northern Adriatic Sea are analysed for the period 1911–1984. An increase in the surface layer and a decrease in the bottom layer are observed in all seasons except winter. Although the oxygen content of the water column as a whole has not changed significantly, it is inferred, from the increasing difference between the surface and the bottom layer, that the primary productivity of the northern Adriatic Sea is increasing. As the average midsummer oxygen content of the bottom layer decreases, the frequency of mass mortality in the benthic fauna is expected to increase, especially in the northern and western subareas.     

Bioconstructional features of the coral Cladocora caespitosa (Anthozoa, Scleractinia) in the Adriatic Sea (Croatia)

The Mediterranean endemic Cladocora caespitosa (Linnaeus, 1767) is a colonial scleractinian coral belonging to the family Faviidae and the only zooxanthellate coral from Mediterranean whose colonies may fuse in reef‐like structures (hermatypic). Recent surveys are focused on three locations where banks occur in the Adriatic Sea (Croatia): near Prvi? Island in the northern Adriatic, near Pag Island in the central Adriatic and in Veliko jezero (Mljet National Park) in the southern Adriatic. The C. caespitosa bank in Veliko jezero covers an area more than 650 m² and is thus the largest bank of C. caespitosa found to date. The strong sea currents, which occur as a result of tidal exchange in the channel, appear to favour the growth of the bank. The goal of the study was to present the influence of major environmental factors upon the build‐up process of the coral bank. Biometrical parameters in the C. caespitosa colonies like diameter of the calyces, polyp ash free dry weight (AFDW), corallite linear growth rate and index of sphericity were investigated and compared from these three locations. The morphology of coral banks from the Adriatic Sea and the disposition of the biometrical values are affected by the sea currents, temperature and sedimentation.     

Sponge disease in the Adriatic Sea

A mass mortality episode involving three marine dyctioceratid sponges, Ircina variabilis, Sarcotragus spinosulus and Spongia officinalis, was observed on ‘Scoglio del Trave’ (Conero Promontory, North Adriatic Sea) in late summer 2009. In this area, calm sea and high temperatures throughout summer created unfavorable environmental conditions, leading to the outbreak of the disease. Affected specimens showed evident necrotic areas and portions with bare skeleton, and many specimens of S. spinosulus were covered with a white mat of cyanobacteria. In September 2009, about 22% of these demosponges suffered from this disease. The mortality event ceased when sea temperature dropped below 20 °C. Small specimens turned out to be more sensitive to the diseases, whereas the damaged tissues of large specimens were able to recover. In October, the damaged specimens were detached by the rough sea and more than 900 beached sponges were found along the 1‐km‐long beach, showing the seriousness of the phenomenon, which probably involved not only the area studied but also areas adjacent to it.     

Contemporary condition of macrozoobenthos in ultrashallow zone of Taman Bay,Sea of Azov

The distribution of macrozoobenthic communities was studied in a vast ultrashallow (0–1.2 m deep) zone of northwestern Taman Bay (separated from the Kerch Strait by Chushka spit) in 2008–2009. Fifty-two species of benthic invertebrates were recorded. The species number, as well as Shannon and Pielou diversity indices, increased along Chushka spit from base to tip. The usual inhabitants of lagoons and estuaries of the Mediterranean Basin and the open Sea of Azov dominated in benthic communities: mollusks Abra segmentum and Hydrobia acuta and polychaetes Heteromastus filiformis and Hediste diversicolor (the latter only in summer). Changes in the community structure were largely determined by the seasonal dynamics of dominant species populations, which was similar to their dynamics in certainother transitional water bodies of the Mediterranean Basin. These changes indicate normal running of seasonal processes in the macrobenthic communities of Taman Bay in 2008 rather than the consequences of a catastrophic black oil spill in the Kerch Strait in November 2007.     

Adriatic ‘opisthobranchs’ (Gastropoda,Heterobranchia): shedding light on biodiversity issues

We herein review the Adriatic opisthobranch fauna, provide an updated checklist of 223 species and assess their distribution at regional and country levels. New Adriatic records are provided for 67 opisthobranch taxa, adding three new records for the Italian coastline, five new records for Albania, eight for Croatia and 15 for Montenegro. The presence of Hermaea bifida (Montagu, 1815), Hermaea variopicta (Costa A., 1869) and Facelina annulicornis (Chamisso & Eysenhardt, 1821) is reported for the first time from anywhere in the Adriatic Sea. Including the new findings, our inventory includes 28 species from Albania, 163 from Croatia, 178 from the Italian coastline of the Adriatic, 41 from Montenegro and 74 from Slovenia. No records were available from Bosnia and Hercegovina. Ninety species (40.4%) are widespread, and were recorded from all three main divisions (Northern, Western and Eastern Adriatic), whilst 79 species (35.4%) were reported from only one of them. At sub‐division levels, the Albanian inventory is the most dissimilar to the other country/regional lists, presumably because of the lack of targeted field surveys. The highest similarity is observed amongst the Western Adriatic, Croatia and the Italian Ionian coastline. Seven alien, one possible alien and three cryptogenic opisthobranchs were reliably recorded from the Adriatic Sea. The first Mediterranean records of five species were in the Adriatic Sea, of which four species have not been found elsewhere in the Mediterranean. Shipping and aquaculture are the probable pathways for most of the alien species. A steady rate of one to two alien introductions per decade since the 1970s was evident. The Northern Adriatic coasts are the most affected by alien invasions, probably due to repeated local introductions along with shellfish transfers (oysters, mussels, Manila clams), and heavy maritime transport.     

Assessing the ecological status of the North-western Adriatic Sea within the European Water Framework Directive: a comparison of Bentix, AMBI and M-AMBI methods

In this paper we looked for the most suitable method for benthic ecological quality status (EcoQS) assessment of the North-western Adriatic Sea. Taking into account the historical and present information, we inter-compared our expectations in terms of EcoQS, with the Bentix, AMBI and M-AMBI classification metrics results, and investigated the sensitivity of these indices to the spatial, seasonal and inter-annual variability of benthic communities. The sampling sites were located at increasing distances with respect to the Po River mouth, along an (inverse) gradient of eutrophication. Moreover, a 2-year monthly monitoring plan (October 2004–September 2006) was carried out at the CEBEB site, located in a central position with respect to the investigated area. The results highlighted that the three indices differed in the sensitivity to spatial and temporal variability of the benthic communities. The strong seasonal and inter-annual changes in the benthic communities had a significant effect on the Bentix, whereas no relevant modifications were observed on the EcoQS assignation obtained using the M-AMBI and AMBI methods. The environmental quality ratios (EQR) values and EcoQS classification obtained by the Bentix and AMBI did not match expectations, as they were not inversely correlated with the site distances with respect to the Po mouth, whereas the M-AMBI clearly discriminated the most and the least eutrophicated sites. It thus appears that the M-AMBI could be considered to be the most valuable method for the ecological-status assessment of the North-western Adriatic Sea benthic ecosystem.     

Spatial segregation of two common Gobius species (Teleostei: Gobiidae) in the northern Adriatic Sea

The habitat characteristics and spatial segregation of the two common gobies Gobius fallax Sarato, 1889 and Gobius auratus Risso, 1810 were investigated by fish frequency and abundance estimates through visual counts in the northern Adriatic Sea. The latter species is represented by a recently described, unusual colour morph in the northern Adriatic Sea. Gobius fallax was observed at five of the nine locations examined and its abundance did not differ between two of three transect locations at Piran (Slovenia) and at the western coast of Cres (Kvarner region, Croatia). Gobius auratus was not found in the Gulf of Trieste but was present at several Kvarner locations, and co‐occurred with G. fallax at one location at Cres. Both inhabit rocky substrates but they show different preferences for substrate inclinations and bathymetric segregation. Gobius fallax was most frequent between 2 and 8 m depth and was most abundant on substrate inclining less than 30°. Its highest abundance over entire transects was 0.3 indiv.·m^?2 but within certain transect sections this increased to 1.25 indiv.·m^?2. Gobius auratus attained its highest abundance in depths below 8 m and at substrate inclinations between 30 and 90°. This species attained its highest values at the western coast of Cres, where it reached a maximum of 1.5 indiv.·m^?2 over entire transects but even up to 3.0 indiv.·m^?2 in certain transect sections. At Krk, the maximum abundance over entire transects was only 0.3 indiv.·m^?2.     

The role of reduced sulfur species in the coalescence of polysaccharides in the Adriatic Sea

Massive mucilage events occur in the Northern Adriatic Sea presumably by cytoplasmic excretions from deteriorated diatoms. During three such events in the summer of 1991, 1997 and 1998 the presence of reduced sulfur species (RSS) was determined in samples of macroaggregate using electrochemical methods (in-phase alternating current (AC) and linear sweep voltammetry). The detected levels of sulfur, expressed as equivalent to sulfide concentrations, were about 200 nM. In the same mucilage samples, concentrations of organic matter were determined in the range from 60 to 600 mg/l of total organic carbon (TOC). The physico-chemical properties of organic matter in the macroaggregates correspond to those of polysaccharides of very high molecular mass. Scanning confocal laser microscopy (SCLM) and fluorescent molecular probes for sugars (the lectins concanvaline A (Con-A)) showed the transformation of polysaccharide polymer structure resulting in the formation of very stable filaments and layers after the treatment of mucilage samples with sodium sulfide. Commercial polysaccharides of bacterial and algal origin (xanthan, carrageenans types I and II, dextran-T-500) have been used to simulate macroaggregate formation under laboratory conditions after treatment with sodium sulfide. Raman spectroscopy indicated that for all model polysaccharides used, sulfide interaction occurred, as evidenced by visible change of the O–H stretching region in the vibration spectra of the water molecules.Our data suggest that the aggregated polysaccharides from the Adriatic Sea are: (1) structurally affected by addition of sulfide as was the case for most of the model polysaccharides, and (2) the stabilizing effect of sulfide on the aggregated polysaccharides is due to the formation of sulfur-organic compounds.     

Mass mortality of the fan mussel,Pinna nobilis in Turkey (eastern Mediterranean)

This study reported the mass mortality events (MMEs) of Pinna nobilis based on diving surveys in Turkey. Data were collected across 12 sites in the Aegean Sea, and one site in the Sea of Marmara, within the period of June-September 2019. The results showed that a low mortality rate of 10% was found in the Sea of Marmara, and a MME was not detected for this area. In contrast, 97% mortality was recorded for the P. nobilis population distributed in the Turkish Aegean Sea. This discrepancy might be related to the seawater temperature and salinity values of these two seas. The waters of the Sea of Marmara were colder and less salty than the Aegean Sea. The findings highlighted the importance of protecting the healthy population of critically endangered fan mussels in the Sea of Marmara. It was recommended that sound management plans comprising of monitoring, enforcement and public awareness activities could be implemented in order to achieve this.     

Freshwater Phytoplankton Bloom in Visovac Lake - A Possible Cause of Benthic Mortality in Krka Estuary (Adriatic Sea, Croatia)

Abstract. In late summer and early autumn of 1989 benthic mortality in the upper part of Krka estuary extended from the Skradinski buk waterfalls to approximately 5 km downstream. Dead benthic organisms were found on the bottom at depths of up to 4.5 m. During August, a freshwater phytoplankton bloom was recorded in Visovac lake (which supplies the Skradinski buk waterfalls). The green tide in the lake was composed of up to 11.9 million cells. 1^--¹, predominantly of Synedra acus and Dinobtyon sertularia . At that time the maximal total inflow of phytoplankton cells through the waterfalls was approx. 0.41 S^-I (38000 1. d^-1). Cells of these two species died after reaching the salt wedge and the cummulative effect of their decomposition apparently caused an oxygen demand sufficient enough to trigger the benthic mortality. Due to the water mass dynamics, the spreading of mortality downstream was slow (from August to October, approx. 5 km). Certain benthic fishes (like Gobius sp.) were observed on the bottom of the affected area in mid-October 1989.     

Annual variability of planktonic ostracods (Crustacea) in the South Adriatic Sea

Temporal variability and population structure of planktonic ostracods were investigated for the first time in the South Adriatic Sea during 1996. The maximal total ostracod abundance (1167 ind·100 m⁻³; 69% juveniles, 18% females and 13% males) was recorded in February. Thirteen species of marine planktonic ostracods were identified. Porroecia spinirostris and Archiconchoecia striata dominated the ostracod assemblage, accounting respectively for 62% and 18% of the total abundance. Their annual peaks were recorded during the cold season, which was the period of their intense reproduction, with favourable temperature conditions and lack of predators. The females surpassed the males in abundance in most species. The presence of the mesopelagic species indicated a strong influence of intermediate layer water masses from the Eastern Mediterranean Sea.     

渤海经济无脊椎动物群落的年代变动

With the decline of fish stocks, the proportions of economically important invertebrates like crustaceans and cephalopods have increased in the Bohai Sea. The community structure and trophic level of economically important invertebrates were analyzed using the bottom trawl survey data collected by the Yellow Sea Fisheries Research Institute in the Bohai Sea in May and August of 1958–1959, 1982, 1992–1993, 2004, 2009 and 2015. A total of 37 species of economically important invertebrates, belonging to 5 orders, 24 families, were captured. The biomass densities of economically important invertebrates in the Bohai Sea displayed an overall downward trend from 1982 to 2015. Oratosquilla oratoria and Loligo spp. were the most dominant species in the past 30 years, the biomass proportion of O. oratoria increased gradually in both May and August from 1982 to 2015. Moreover,biodiversity indices of economically important invertebrates in the Bohai Sea appeared to decline from 1982 to2004 and then increased in 2015. Similarly, the mean trophic level of economically important invertebrates declined from 1982 to 2004 and increased slightly in 2015. Overall, although the proportions of invertebrates have increased, the biomass densities in the Bohai Sea have displayed an overall downward trend from 1982 to 2015.The increases in the biodiversity and trophic level of economically important invertebrates after the 2000 s,possibly benefit from stock enhancement projects implemented by governments at different levels and national fishery management measures such as the "double-control" of the total number and engine power of fishing vessels and summer moratorium of fishing.     

Benthic assemblage composition on subtidal reefs along a latitudinal gradient in Western Australia

At regional scales, the distribution of species and the structure of assemblages vary with latitude within many marine and terrestrial systems. The oligotrophic coastal waters of Western Australia (WA) support highly speciose and endemic assemblages, yet spatial patterns in benthic structure are currently poorly known. We examined benthic assemblage composition along a latitudinal gradient of 28.5–33.5°S and a depth gradient of 14–62 m, on subtidal reefs in warm-temperate WA. We surveyed benthos using a remotely triggered digital stills camera. In total, we sampled macroalgae and sessile invertebrates at 201 sites spread across four locations. Percent cover of coarse taxonomic groups and dominant species was estimated from over 2000 photoquadrat samples. We recorded significant differences in benthic assemblage composition between locations, and along depth gradients within each location. However, the magnitude of change with depth was not consistent between locations, and shifts in assemblage composition along the depth gradients were not as pronounced as expected. The percent cover of all dominant benthic groupings differed between locations, and several key taxa, such as the kelp Scytothalia dorycarpa, brown foliose macroalgae, hard corals and sponges, changed predictably along the latitudinal gradient. Our study adopted a coarse taxonomic, but assemblage-wide, approach to describing macrobenthic assemblages, and clear differences between locations and depths were detected. The surveys have provided baseline data on broad scale ecosystem structure against which to detect future ecological change.     

Distribution of Sessile Invertebrates in a Rocky Sublittoral Community off Rimsky‐Korsakov Islands (Sea of Japan)

Abstract. The dependence of the vertical distribution of sessile macrobenthos upon environmental physical factors was studied in a community inhabiting rocky substrata at 2 to 15 m depth near the Rimsky‐Korsakov Islands (Peter the Great Bay, Sea of Japan). Three separate groups of species were distinguished within this community. The main dominant species are the anthozoan Metridium senile and the mussel Crenomytilus grayanus. Water movement, bottom inclination and topographic position of the studied sites in respect to prevailing water flows influence the spatial structure of sessile invertebrates. No vertical zonation in the scale of the studied sublittoral area below the 2 m depth level was observed. The studied factors are believed to condition the observed mosaic of patches characterized by specific composition, with a single species being dominant.     

The large deep water transient in the Eastern Mediterranean

The recent changes in the thermohaline circulation of the Eastern Mediteranean caused by a transition from a system with a single source of deep water in the Adriatic to one with an additional source in the Aegean are described and assessed in detail. The name Cretan Sea Overflow Water (CSOW) is proposed for the new deep water mass. CSOW is warmer (θ>13.6°C) and more saline (S>38.80) than the previously dominating Eastern Mediterranean Deep Water (EMDW), causing temperatures and salinities to rise towards the bottom. All major water masses of the Eastern Mediterranean, including the Levantine Intermediate Water (LIW), have been strongly affected by the change. The stronger inflow into the bottom layer caused by the discharge of CSOW into the Ionian and Levantine Basins induced compensatory flows further up in the water column, affecting the circulation at intermediate depth. In the northeastern Ionian Sea the saline intermediate layer consisting of Levantine Intermediate Water and Cretan Intermediate Water (CIW) is found to be less pronounced. The layer thickness has been reduced by factor of about two, concurrently with a reduction of the maximum salinity, reducing advection of saline waters into the Adriatic. As a consequence, a salinity decrease is observed in the Adriatic Deep Water. Outside the Aegean the upwelling of mid-depth waters reaches depths shallow enough so that these waters are advected into the Aegean and form a mid-depth salinity-minimum layer. Notable changes have been found in the nutrient distributions. On the basin-scale the nutrient levels in the upper water column have been elevated by the uplifting of nutrient-rich deeper waters. Nutrient-rich water is now found closer to the euphotic zone than previously, which might induce enhanced biological activity. The observed salinity redistribution, i.e. decreasing values in the upper 500–1400 m and increasing values in the bottom layer, suggests that at least part of the transition is due to an internal redistribution of salt. An initiation of the event by a local enhancement of salinity in the Aegean through a strong change in the fresh water flux is conceivable and is supported by observations.