On three interesting marine red algae (Ceramiales,Rhodophyta) from the Mediterranean Sea

Abstract

Observations on three interesting Ceramiales from the Mediterranean Sea are reported; Ceramium incospicuum Zanardini, Polysiphonia setacea Hollenberg and Rodriguezella pinnata (Kützing) Schmitz ex Falkenberg. The male reproductive structures on the genus Rodriguezella are described for the first time.     

The reproductive structures of the Mediterranean alga Laurencia pelagosae (Ceramiales,Rhodophyta)

Reproductive structures of Laurencia pelagosae are described for the first time. Tetrasporangia, in parallel arrangement, are cut off from the mother cells adaxially; antheridial branches, which are unramified, are inserted in depressions lacking a row of axial cells on their bases; cystocarps are sessile and subspherical. The occurrence of secondary pit-connections between epidermal cells and of lenticular thickenings in the walls of medullary cells is documented. The species is included in the new section Pelagosae within the subgenus Laurencia.     

The occurrence of Gonyaulax tamarensis Lebour bloom in the Adriatic Sea along the coast of Emilia-Romagna

Abstract

The authors describe the occurrence of Gonyaulax tamarensis Lebour bloom in the Adriatic Sea between 200 and 1000 m off the coast of Emilia-Romagna during the period of August 14-20 1982.     

Orientation of crustacean burrows in the Bay of Panzano (Gulf of Trieste, Northern Adriatic Sea)

Three burrow systems produced by Jaxea nocturna and one cluster of burrows produced by Upogebia pusilla where investigated in the Bay of Panzano, Northern Adriatic Sea, to determine preferred orientations. The distributions of dip directions differ between both producers. Steep shafts into the consolidated mud followed by large, shallowly inclined tunnels oriented in a manner similar to a spiral are characteristic for Jaxea nocturna burrows. In contrast, the Y-shaped burrows of Upogebia pusilla have entrance shafts that are less steep which are connected by a near-horizontal section, where a blind ending tunnel branches. Orientations of the dominating, shallowly inclined burrow parts are not randomly distributed in either Jaxea or in Upogebia burrows. Three preferred axial orientations with almost equal proportions in each direction are characteristic for the Jaxea burrow from the centre of the bay. This system transforms to a squared structure near the coast, where one direction parallels the shoreline and the other is oriented at right angles to the coast. The latter orientation demonstrates significant constancy in all investigated burrows. Upogebia burrow clusters coincide with the three preferred orientations of the Jaxea burrow from the bay centre, but the proportions of the directions are unequal. Burrow segments connecting the steep entrance shafts in Upogebia are oriented almost parallel to the shoreline, whereas at right angles to the coast the flat blind-ending tunnels incline towards the open sea. The study shows a strong coincidence between empirical and theoretical distributions of the dominating orientations in both species.     

Orientation of crustacean burrows in the Bay of Panzano (Gulf of Trieste, Northern Adriatic Sea)

Three burrow systems produced by Jaxea nocturna and one cluster of burrows produced by Upogebia pusilla where investigated in the Bay of Panzano, Northern Adriatic Sea, to determine preferred orientations. The distributions of dip directions differ between both producers. Steep shafts into the consolidated mud followed by large, shallowly inclined tunnels oriented in a manner similar to a spiral are characteristic for Jaxea nocturna burrows. In contrast, the Y-shaped burrows of Upogebia pusilla have entrance shafts that are less steep which are connected by a near-horizontal section, where a blind ending tunnel branches. Orientations of the dominating, shallowly inclined burrow parts are not randomly distributed in either Jaxea or in Upogebia burrows. Three preferred axial orientations with almost equal proportions in each direction are characteristic for the Jaxea burrow from the centre of the bay. This system transforms to a squared structure near the coast, where one direction parallels the shoreline and the other is oriented at right angles to the coast. The latter orientation demonstrates significant constancy in all investigated burrows. Upogebia burrow clusters coincide with the three preferred orientations of the Jaxea burrow from the bay centre, but the proportions of the directions are unequal. Burrow segments connecting the steep entrance shafts in Upogebia are oriented almost parallel to the shoreline, whereas at right angles to the coast the flat blind-ending tunnels incline towards the open sea. The study shows a strong coincidence between empirical and theoretical distributions of the dominating orientations in both species.     

Ultrastructure of post-fertilization development in the red alga Nienburgia andersoniana (Delesseriaceae, Ceramiales, Rhodophyta)

The ultrastructure of post-fertilization development in Nienburgia andersoniana (J. Ag.) Kyl. is described. Above the auxiliary cell there is a group of four sterile cells. The presence of abundant storage products (starch granules, lipid bodies and protein crystals) in these cells indicates that the sterile cells function as nutrient suppliers to the young auxiliary and gonimoblast cells of the carposporophyte during its early steps of development. Following fertilization and transfer of the diploid nucleus to the auxiliary cell, the trichogyne disappears and large multinucleate gonimoblast initials are produced. These subsequently produce generative gonimoblast cells which cleave successively to form young carpospores. Those of the gonimoblast cells which will not differentiate into carpospores are transformed into cells producing mucilage. Both kinds of gonimoblast cells contain plastids, starch granules, cytoplasmic concentric membrane bodies and small vesicles. Dark-staining spherical masses occurring in the cytoplasm of the auxiliary and gonimoblast cells may represent degenerating haploid nuclei. Septal plugs interconnecting the auxiliary cell and gonimoblast cells increase considerably in size during carposporophyte development. The fusion cell at the late stage of carposporophyte development appears degenerative. Young carpospores have plastids and mitochondria, and concentric membrane bodies that will form mucilage sacs. Medium-aged carpospores have fully developed plastids, starch granules and fibrous vacuoles. Mature carpospores possess, in addition, cored vesicles. The inner pericarp cells contribute large amounts of mucilage to the cytostocarpic cavity and eventually are consumed. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 289–299.     

Polysiphonia perforans sp. nova (Ceramiales,Rhodophyta) from the Mediterranean Sea

ABSTRACT

A new species of the genus Polysiphonia from the Mediterranean Sea is described. It is an ecorticate species, with 4 pericentral cells, showing prostrate axes from which erect axes arise. Rhizoids are formed by pericentral cells in a median position remaining in open connection with them. Erect axes are straight, simple throughout or pseudodichotomous at the base then simple or with one (rarely two-three) orders of branching; trichoblasts and scar cells not observed. Only tetrasporangial plants were found. They show ellipsoid tetrasporangia borne in short straight series, often interrupted by sterile segments. The new species is also characterized by a peculiar habit consisting of prostrate axes, adhering to lower faces of Peyssonnelia spp. by means of rhizoids growing upward, from which erect upright axes perforating thalli of the supporting species arise. A comparison with the related species of Polysiphonia was also carried out.     

Ceramium gobii n. sp. (Rhodophyta, Ceramiales), a brackish-water red alga in the Baltic Sea

A geographical survey was made of the distribution of diaphanoid Ceramia in the Baltic Sea and adjacent waters west of the Baltic. It was found that Ceramium plants with so-called 'Gobi's' parasporangia (monosporangia) have a wide distribution in the Baltic Sea at salinities of ca (3) 5–8%. This brackish-water Ceramium is here described as a new species, Ceramium gobii. At higher salinities in the south of the Baltic Sea and on the Swedish west coast, round parasporangia (polysporangia) indicate the occurrence of other Ceramium species. The new species is very similar to the brackish-water species Ceramium radiculosum , which was described from river-mouths to the Gulf of Trieste (Italy). A comparison was made of several diaphanoid Ceramium species, C. diaphanum, C. tenuicorne, C. gobii , and C. radiculosum , including (1) size of male plants, (2) type of paraspores, (3) vegetative characters.     

The benthic algal flora on rocky substrata of the Tremiti Islands (Adriatic Sea)

ABSTRACT

The results of a floristic study of benthic marine macroalgae on rocky substrata from the Tremiti Islands (Adriatic Sea) are presented. The list of taxa at specific and infraspecific level consists of 226 Rhodophyceae, 59 Fucophyceae and 36 Chlorophyceae. Of these 321 taxa, 147 are newly reported from the Tremiti Islands. Data on the reproductive phenology of each species, and comments on some species of particular interest are also given.     

Ultrastructure of auxiliary and gonimoblast cells during carposporophyte development in the red alga Cryptopleura ruprechtiana (Delesseriaceae, Ceramiales, Rhodophyta)

The ultrastructure sequence for the complete post-fertilization development is described in Cryptopleura ruprechtiana (C. Agardh) Kylin, a member of the Delesseriaceae. Following fertilization the diploid nucleus is transferred to the auxiliary cell. This contains typical red algal proplastids, cytoplasmic concentric membranes, numerous small vacuoles and lipid bodies. Crystalline inclusions and virus-like particles are also present. In addition darkly staining spherical masses possibly represent dehydrated haploid chromatin. The multinucleate auxiliary cell produces initially one large gonimoblast initial and subsequently many smaller gonimoblast initials. The first formed generative gonimoblast cell is similar in cellular structure to the auxiliary cell. Gonimoblast initials are uninucleate but through caryokinesis they become multinucleate. They undergo repeated cleavage to form more gonimoblast cells. Subsequent, centripetal cytokinesis results in the formation of clusters of gonimoblast cells. A new type structural cap or association is observed in the septal plugs that interconnect gonimoblast initials. Terminal or generative gonimoblast cells cleave to form additional gonimoblast cells. Only terminal gonimoblast cells are differentiated to carpospores.     

Zooplankton community and hydrographical properties of the Neretva Channel (eastern Adriatic Sea)

Temporal and spatial variability of micro and mesozooplankton was studied in 1998 and 1999 at four stations in the Neretva Channel area influenced by the Neretva river and the open waters of the south Adriatic Sea. The area is orthophosphate limited, but an excessive accumulation of land derived nitrogen is prevented by phytoplankton uptake and the general circulation pattern. Microzooplankton was dominated by ciliates, with average abundances comparable to other Adriatic channel areas (122–543 ind. l⁻¹). Non-loricate ciliates (NLC) generally peaked in the warmer periods, but a winter increase was evident towards the inner part of the channel. Tintinnid abundances generally increased in autumn. A significant relationship with temperature was not recorded for either protozoan group. An inverse relationship between NLC and salinity might be indirectly caused by their preference for the food abundant surface layer. Mesozooplankton was dominated by copepods, with distinct summer maxima throughout the area and pronounced winter maxima of >10,000 ind. m⁻³ at the inner stations. The community was predominantly neritic but the open sea waters were important in structuring the mesozooplankton assemblage at all stations during the autumn–winter period. Although temperature regulated the seasonal dynamics of most metazoans and the species succession in the copepod community, small omnivorous copepods (Oncaea media complex, Oithona nana and Euterpina acutifrons) dominated regardless of the season. A trophic link between copepods and ciliates was evident in winter during low phytoplankton biomass.     

Assessment of nutrient limitation in Rijeka Bay, NE Adriatic Sea, using miniaturized bioassay

In this paper we consider the trophic state and ambient nutrient limitation by the trophic index (TRIX) calculation and nutrient enrichment bioassay using the diatom Phaeodactylum tricornutum at two differently eutrophicated NE coastal Adriatic locations (in the Rijeka Bay area). The first station in Rijeka Port had significantly higher concentrations of dissolved inorganic nutrients and TRIX indices (up to 7.53) than the station near Opatija. The DIN/TP ratios in Rijeka Port were lower (11.03 to 44.47) than those in Opatija (38.67 to 82.00), indicating P limitation in Opatija. The nutrient enrichment bioassays revealed P as the key limiting nutrient for the diatom growth at both sites. In addition, silica limitation was found in all water samples with the exception of the surface water samples in Rijeka Port. Nitrogen limitation was rarely evident.     

Ecology and taxonomy of potentially toxic Pseudo-nitzschia species in Lim Bay (north-eastern Adriatic Sea)

The population dynamics of Pseudo-nitzschia in relation to environmental factors was investigated from March 2002 to July 2008 in Lim Bay, in the north-eastern Adriatic Sea. Domoic acid was monitored in the breeding population of Mytilus galloprovincialis from 2005 to 2008. The principal-component analysis of environmental parameters showed that the system is mostly temperature driven. The phytoplankton community was mainly composed of diatoms. Pseudo-nitzschia was the dominant diatom, present in 60% of samples, with a maximum (1.6 × 10⁶ cells L⁻¹) contribution up to 97% of the total diatom abundance. Morphological analysis revealed Pseudo-nitzschia manii and potentially toxic Pseudo-nitzschia pseudodelicatissima, Pseudo-nitzschia pungens, Pseudo-nitzschia fraudulenta and Pseudo-nitzschia calliantha as the dominant species in blooms. Pseudo-nitzschia abundance positively correlated to temperature, phosphate and ammonia in accordance with its maximal abundance in the summer/autumn period when fish farms had a maximum impact on the environment. Domoic acid was detected in M. galloprovincialis in concentrations below regulatory limits, ranging from 0.097 to 0.8721 μg g⁻¹ in five cases from April to October 2005 in Lim Bay, but so far it is not clear which of the species was responsible for DA production. This study is also the first record of P. manii, P pungens and P. fraudulenta species in the Adriatic Sea.     

The evolution and phytoplankton composition of mucilaginous aggregates in the northern Adriatic Sea

Recurrent occurrences of visible mucilage “clouds” that cover areas up to several hundred kilometres with vertical dimensions of 20–30 m have been recorded in the stratified water column in the northern Adriatic. In the past this was described as “mare sporco” phenomenon. Past studies of the phenomenon indicated that phytoplankton is an important component of mucilage. Our research revealed the composition of phytoplankton assemblages in different types of mucilaginous aggregates collected during the summers of 1997 and 2000 using pigment biomarkers (HPLC). Phytoplankton biomass in the mucilage samples was very high, ranging from 7.9 μg g⁻¹ to 390.8 μg g⁻¹ of chlorophyll a per unit of dry mass of mucilage. The phytoplankton community in the early, loose stage of mucilaginous aggregates was heterogeneous, as indicated by the diversity of detected pigments. The number of phytoplankton groups decreased as the aggregates aged and diatoms increased in relative biomass (up to 92.7%). Phytoplankton biomass in seawater was similar in years with and without mucilage; however, significantly higher contributions to the total biomass of 19′-hexanoyloxyfucoxanthin-containing phytoplankton (prymnesiophytes) were found in the upper 10 m in spring of the “mucilaginous years” (1997 and 2000) followed by prevalence of diatoms in summer. The F_p pigment index used to assess seawater trophic conditions reached lower values in April–May in mucilaginous years (1997 and 2000) compared to non-mucilaginous years (1998 and 1999). We conclude that the role of prymnesiophytes and other small flagellates is crucial for the initial phases of mucilage appearance. Aggregates represent a favourable environment for the secondary development of opportunistic diatoms that foster mucilage formation.     

The Colonisation of Human-made Structures by the Invasive Alga Codium fragile ssp. tomentosoides in the North Adriatic Sea (NE Mediterranean)

Human-made structures, such as groynes, breakwaters, seawalls, pier pilings and floating pontoons, are becoming common features of the landscape in urbanised coastal and estuarine areas. Despite this tendency few studies have focused on their ecology or on their potential impacts on natural assemblages of organisms. When artificial structures are introduced in areas with little or no hard substrata, they not only provide novel habitats, which enables the colonisation of sandy areas by hard-bottom dwelling species, but they can also provide suitable habitats for exotic species. Along the north-east coast of Italy, sandy shores are protected from erosion by a line of breakwaters, which runs almost uninterrupted for about 300 km. These structures provide habitat for a variety of macroalgae and invertebrates and also for the invasive green alga Codium fragile ssp. tomentosoides. The aim of this study was, therefore, to investigate patterns of distribution of this alga on breakwaters in Cesenatico. In particular, we compared the density of thalli, biomass, length and degree of branching of C. fragile ssp. tomentosoides between the landward and the seaward sides of breakwaters, to test the hypothesis that sheltered habitats (landward) represent more suitable habitats than exposed habitats (seaward). In general, the landward side of breakwaters supported greater numbers of thalli of C. fragile ssp. tomentosoides than seaward sides. Thalli grew longer and more branched in sheltered habitats, leading to an overall larger biomass of the alga on the landward side of breakwaters. The presence of sheltered human-made hard substrata in the vicinity of major trading ports and sources of eutrophication could enhance the dispersal of invasive species across regional and geographic scales. Thus, the effects of artificial structures and introduced species on coastal assemblages cannot be evaluated separately, but their synergistic nature should be considered in planning strategies for conservation of biodiversity in coastal habitats.     

Patterns of genetic diversity of the cryptogenic red alga Polysiphonia morrowii (Ceramiales,Rhodophyta) suggest multiple origins of the Atlantic populations

The red alga Polysiphonia morrowii, native to the North Pacific (Northeast Asia), has recently been reported worldwide. To determine the origin of the French and Argentine populations of this introduced species, we compared samples from these two areas with samples collected in Korea and at Hakodate, Japan, the type locality of the species. Combined analyses of chloroplastic (rbcL) and mitochondrial (cox1) DNA revealed that the French and Argentine populations are closely related and differ substantially from the Korean and Japanese populations. The genetic structure of P. morrowii populations from South Atlantic and North Atlantic, which showed high haplotype diversity compared with populations from the North Pacific, suggested the occurrence of multiple introduction events from areas outside of the so‐called native regions. Although similar, the French and Argentine populations are not genetically identical. Thus, the genetic structure of these two introduced areas may have been modified by cryptic and recurrent introduction events directly from Asia or from other introduced areas that act as introduction relays. In addition, the large number of private cytoplasmic types identified in the two introduced regions strongly suggests that local populations of P. morrowii existed before the recent detection of these invasions. Our results suggest that the most likely scenario is that the source population(s) of the French and Argentine populations was not located only in the North Pacific and/or that P. morrowii is a cryptogenic species.     

The cell wall chemistry of Bangia atropurpurea (Bangiales, Rhodophyta) and Bostrychia moritziana (Ceramiales, Rhodophyta) from marine and freshwater environments

The cell wall polysaccharides of two species of red algae, which are adapted to both freshwater and marine environments, were analysed to determine the effect of these widely different environments on their commercially important agarocolloids and to investigate the possible role of the cell wall in environmental adaptation. Cell wall polymers of freshwater isolates of Bangia atropurpurea (Roth) C. Agardh and cultured freshwater and marine Bostrychia moritziana (Sonder ex Kützing) J. Agardh were isolated and the polysaccharides chemically fractionated and characterized. Wall polysaccharides of freshwater B. atropurpurea were similar to those previously reported for marine isolates with repeating disac-charide units of agarose and porphyran predominant in the hot water extracts. In the insoluble residues, 3-iinked galactosyl and 4-linked mannosyl residues were predominant. Bostrychia moritziana wall polysaccharides included agarocolloids with various patterns of methyl ether substitution similar to those previously described for other Ceramiales. Differences in the position of methyl ether substituents were detected in the hot water extracts of the freshwater and marine specimens. Polymers of freshwater ß. moritziana cultures were composed of a complex mixture of repeating disaccharide units including 2′-O-methyl agarose, 6-O-methyI agarose and 2′-O-methyl porphyran. Polymers of marine isolates of ß. moritziana differ in that they contain only trace amounts of 2-O-methyl saccharides and increased amounts of 6-O-amethyl saccharides. The hot water insoluble residues of both freshwater and marine isolates of ß. moritziana contain a mixture of 3-linked galactosyl and 4-linked glucosyl residues. These results indicate that the adaptive response of B. moritziana to changing osmotic and ionic conditions may include changes in cell wall chemistry: notably, the pattern of methyl ether substitution.     

Genetic diversity of the mangrove‐associated alga Bostrychia radicans/Bostrychia moritziana (Ceramiales,Rhodophyta) from southern Central America

Previous studies suggested that the biodiversity of the mangrove‐associated Bostrychia radicans/Bostrychia moritziana species complex on the Pacific coast of Central America, based on genetic and reproductive data, were low compared with similar areas on the Atlantic coast. Evolutionary scenarios were proposed based on either a recent introduction to the Pacific, or a more uniform environment leading to genetically connected populations and low differentiation between populations. We sampled more extensively in southern Mexico, Guatemala and El Salvador and sequenced the samples for the RuBisCo spacer. Our results show that genetic diversity is high in these populations. Many haplotypes retrieved are also found in the Atlantic Ocean (USA, Brazil), an observation not made before. Data suggest that populations are highly differentiated with little evidence of isolation‐by‐distance. The population at La Puntilla, El Salvador is highly differentiated from other populations. Data also suggest that diversity is reduced in a northerly direction, with only one haplotype, unique to Pacific Central America, found north of Chiapas, Mexico. This could be due to northern expansion of this unique genotype as sea surface temperatures ameliorated following the last glacial maximum. Our data do not support the previous proposition of low diversity in the east central Pacific and suggest that much of the Pacific Central America diversity is from before the closure of the Isthmus of Panama.     

Cartography of littoral rocky-shore communities (CARLIT) as a tool for ecological quality assessment of coastal waters in the Eastern Adriatic Sea

Littoral cartography is a valuable tool which uses shallow water macroalgal communities for monitoring ecosystem health and water quality. Cartography of littoral rocky-shore communities (CARLIT) is a European Union Water Framework Directive-compliant monitoring method widely used in the Western Mediterranean Sea. This non-destructive method is based on a visual observation of the type and length of coast occupied by rocky-shore communities in the upper-sublittoral zone. Here we present the first wide-scale application of CARLIT in the Adriatic Sea. The reference values calculated specifically for the Croatian coast (Eastern Adriatic Sea) ensure accurate calculation of the ecological status. The measured ecological quality ratio values (EQR) show a good linear relation with the Land Uses Simplified Index (LUSI) and appropriately reflect an increase in nutrient concentrations. The results suggest that this method can be used as a reliable biomonitoring tool in the Eastern Adriatic Sea. This study highlighted that Cystoseira forests thrive in 39% of the surveyed coastline and 23% of the coastline seems to be subjected to overgrazing by sea urchins. Our results provide a spatially accurate information on the abundance of different upper-sublittoral communities and the associated ecological status, which can be used in future management plans for improving water quality.