Pseudo-nitzschia species population dynamics in the Quoddy Region, Bay of Fundy

The population dynamics of Pseudo-nitzschia species from the Quoddy Region of the Bay of Fundy was examined in a field-based study. Five stations were sampled over a period of 11 weeks during the course of one discrete bloom episode in 2003 with seven species of Pseudo-nitzschia found and enumerated: P. americana, P. delicatissima, P. pseudodelicatissima, P. fraudulenta, P. pungens, P. seriata and P. subpacifica. We related species abundance to physical and chemical properties of seawater (transparency, fluorescence, silicate, phosphate, nitrite + nitrate, ammonia, nitrite, oxygen, sigma-t, tidal level, tidal state and total depth of water column) using canonical correspondence analysis (CCA) to identify factors explaining the greatest amount of variance in their temporal and spatial distribution patterns. Our results indicate that abundance of species and groups of species correlated well with certain specific chemical and physical properties of seawater. P. pseudodelicatissima and P. delicatissima abundance was positively correlated with nitrates and P. americana with depth of the water column. P. pungens was more abundant in samples with higher concentrations of phosphates and lower concentrations of nitrates. P. seriata abundance was negatively associated with total fluorescence. P. subpacifica and P. fraudulenta abundances were not statistically related to any of the variables examined. Our data therefore provides direction for testable, hypothesis driven experiments that could provide predictive insights into the occurrence of certain harmful algal bloom (HAB) species should specific environmental variables be further affected along the gradients extracted in this study.     

Intra- and interspecies differences in growth and toxicity of Pseudo-nitzschia while using different nitrogen sources

Clonal cultures of plankton are widely used in laboratory experiments and have contributed greatly to knowledge of microbial systems. However, many physiological characteristics vary drastically between strains of the same species, calling into question our ability to make ecologically relevant inferences about populations based on studying one or a few strains. This study included 19 non-axenic strains of three species of the diatom Pseudo-nitzschia isolated primarily from the mid-Atlantic coastal region of the United States. Toxin (domoic acid) production and growth rates were measured in cultures using different nitrogen sources (NH₄⁺, NO₃⁻ and urea) and growth irradiances. The strains exhibited broad differences in growth rate and toxin content even between strains isolated from the same water sample. The influence of bacteria on toxin production was not investigated. Both P. multiseries clones produced toxin, yet preferentially used different nitrogen sources. Only two of nine P. calliantha and two of five P. fraudulenta isolates were toxic and domoic acid content varied by orders of magnitude. All three species had variable intraspecies growth rates on each nitrogen source, but P. fraudulenta strains had the broadest range. Light-limited growth rate and maximum growth rate in P. fraudulenta and P. multiseries varied with species. These findings show the importance of defining intra- and interspecies variability in ecophysiology and toxicity. Ecologically relevant functional diversity in the form of ecotypes or cryptic species appears to be present in the genus Pseudo-nitzschia.     

Pseudo-nitzschia and domoic acid fluxes in Santa Barbara Basin (CA) from 1993 to 2008

Blooms of domoic acid (DA) producing Pseudo-nitzschia, regularly occur off the coast of California. Although it has been hypothesized that these blooms are increasing in frequency, the lack of historical records limits our understanding of potential causal mechanisms. In this study, an 15-year time-series (1993–2008) of sediment trap samples collected from the Santa Barbara Basin (SBB) at 540 m were analyzed for Pseudo-nitzschia (n = 196, microscopy and SEM) and DA (n = 206, LC–MS/MS) concentrations and fluxes. Results suggest that there was an abrupt shift towards greater frequency and higher magnitude Pseudo-nitzschia blooms and toxic DA flux events in the SBB after the year 2000. SEM analysis of sediment trap material indicates that these events were mainly blooms of P. australis, with cell fluxes increasing by an order of magnitude from a maximum of 4.5 × 10⁶ cells m⁻² d⁻¹ pre-2000, to as high as 3.2 × 10⁸ cells m⁻² d⁻¹ thereafter. Similarly, sediment trap DA fluxes increased by an average of 13.4 μg m⁻² d⁻¹, with only one large event (>5 μg m⁻² d⁻¹) from 1993 to 1999 versus 16 large DA events from 2000 to 2008. While the causes of this abrupt shift remain ambiguous, we suggest that this shift may be related to natural climate variability associated with a change in phase of the North Pacific Gyre Oscillation (NPGO) and its potential influence on the composition and magnitude of waters that are upwelled into the SBB.     

Note on the occurrence of Pseudo-nitzschia australis and domoic acid in squid from Monterey Bay, CA (USA)

Over the past decade diatom blooms of domoic acid (DA)-producing Pseudo-nitzschia spp. have been responsible for numerous marine mammal and bird mortalities in Monterey Bay, CA. One possible toxin vector is the market squid, Loligo opalescens, a small pelagic mollusk that plays an important role in the near-shore food web of the California Current ecosystem as a favored vertebrate prey species. This study examined the trophic link between toxic Pseudo-nitzschia and L. opalescens using toxin and stomach content analyses of animals collected from Monterey Bay, CA in 2000. Receptor binding assay data (confirmed by tandem mass spectrometry), demonstrated the presence of DA in squid during a toxic Pseudo-nitzschia event, with P. australis frustules observed in stomach samples. Though DA levels were low (<0.5 μg DA g⁻¹ tissue) in L. opalescens during the study period, it is now clear that this potent neurotoxin can occur in squid and is likely delivered through its krill prey species, which are known to retain DA after feeding on toxic Pseudo-nitzschia. Our findings suggest that further study of the relationship between Pseudo-nitzschia blooms and DA contamination of squid is warranted to better evaluate the potential health risk to humans and wildlife associated with this major commercial seafood species and important prey item.     

我国沿海拟菱形藻属的2新记录种及其产毒特征分析

为澄清我国沿海拟菱形藻属(Pseudo-nitzschia)的物种多样性,并确认中国海域拟菱形藻属是否具有产生多莫酸(Domoicacid)的能力,采用毛细管显微操作技术从我国沿海水体中分离、纯化拟菱形藻细胞,建立了单克隆培养株系,并基于核糖体转录间隔区ITS1-5.8S-ITS2 序列构建了分子系统树。结果表明,结合在光学显微镜和透射电镜下观察的形态学特征和分子系统发育分析数据,鉴定到我国拟菱形藻属的2新记录种:银河拟菱形藻(P. galaxiae Lundholm & Moestrup)和微孔拟菱形藻(P. micropora Priisholm, Moestrup & Lundholm),对其形态学特征进行了详细描述,并与相似种类进行了比较研究。利用高效液相色谱(HPLC)技术对多莫酸特征进行了检测,结果表明培养株系并不产生多莫酸。这些为我国拟菱形藻属物种多样性和产毒特征研究提供了基础数据。     

Growth dynamics of non-toxic Pseudo-nitzschia delicatissima and toxic P. seriata (Bacillariophyceae) under simulated spring and summer photoperiods

Marine planktonic diatoms of the genus Pseudo-nitzschia Peragallo have been responsible for amnesic shellfish poisoning (ASP) events worldwide through the production of the neurotoxin domoic acid (DA). The appearance and toxicity of Pseudo-nitzschia species is variable throughout the year and potentially linked to changes in environmental parameters; many ASP events occur in relatively high latitudes where day length is particularly variable with season. In UK waters, shellfish monitoring has prevented any impact on human health but has led to long-term closures of fisheries, with severe economic consequences. Laboratory experiments on two Pseudo-nitzschia species typically found in Scottish West Coast waters during spring (short photoperiod (SP)) and summer (long photoperiod (LP)) conditions were conducted to determine the influence of photoperiod on their growth and toxicity. Results indicated that non-toxic P. delicatissima (Cleve) Heiden achieved a greater cell density under SP (9-h light:15-h dark (L:D) cycle). For toxin-producing P. seriata (Cleve) H. Peragallo, a LP (18-h L:6-h D cycle) resulted in an enhanced growth rate, cell yield and total toxin production, but it decreased the toxin production per cell. A better understanding of the response of Pseudo-nitzschia species to photoperiod and other foreseeable environmental variables may help predict the appearance of toxic strains.     

Correlation between the presence of Gonyaulax fragilis (Dinophyceae) and the mucilage phenomena of the Emilia-Romagna coast (northern Adriatic Sea)

The formation of massive amounts of suspended mucilaginous organic matter which periodically affects the Adriatic Sea, has been regarded as a complex physico-chemical phenomenon resulting from the production of extracellular material by phytoplankton. Although the exact cause has remained obscure, the mechanism of its formation has usually been considered to be a long-term process, starting after the late winter-early spring blooms, and involving the participation of various algal species, mainly within the diatom group. In this paper we report on the results of a phytoplankton monitoring programme in northern Adriatic seawaters off the Emilia-Romagna coast of Italy which revealed the constant concomitant presence of the dinoflagellate Gonyaulax fragilis (Schütt) Kofoid and mucilaginous formations. In the early stages of the phenomenon the dinoflagellate was clearly observable by microscopic examination in the mucilage, but as the mucilage aged this alga almost completely decomposed and diatom cells increased in number and became predominant. Although characterized by a slow growth rate in culture, in natural seawater G. fragilis was observed to reach cell densities of up to 7.0×10⁶ cells l⁻¹. The results of this study lead us to propose the hypothesis that the appearance of mucilage in the water column of the Adriatic Sea is the consequence of a seasonal growth of this dinoflagellate favoured by specific environmental circumstances.     

Blooms of Pseudo-nitzschia and domoic acid in the San Pedro Channel and Los Angeles harbor areas of the Southern California Bight, 2003–2004

Abundances of Pseudo-nitzschia spp. and concentrations of particulate domoic acid (DA) were determined in the Southern California Bight (SCB) along the coasts of Los Angeles and Orange Counties during spring and summer of 2003 and 2004. At least 1500 km² were affected by a toxic event in May/June of 2003 when some of the highest particulate DA concentrations reported for US coastal waters were measured inside the Los Angeles harbor (12.7 μg DA L⁻¹). Particulate DA levels were an order of magnitude lower in spring of 2004 (February and March), but DA concentrations per cell at several sampling stations during 2004 exceeded previously reported maxima for natural populations of Pseudo-nitzschia (mean = 24 pg DA cell⁻¹, range = 0–117 pg DA cell⁻¹). Pseudo-nitzschia australis dominated the Pseudo-nitzschia assemblage in spring 2004. Overall, DA-poisoning was implicated in >1400 mammal stranding incidents within the SCB during 2003 and 2004. Ancillary physical and chemical data obtained during our regional surveys in 2004 revealed that Pseudo-nitzschia abundances, particulate DA and cellular DA concentrations were inversely correlated with concentrations of silicic acid, nitrogen and phosphate, and to specific nutrient ratios. Particulate DA was detected in sediment traps deployed at 550 and 800 m depth during spring of 2004 (0.29–7.6 μg DA (g sediment dry weight)⁻¹). The highest DA concentration in the traps was measured within 1 week of dramatic decreases in the abundances of Pseudo-nitzschia in surface waters. To our knowledge these are the deepest sediment trap collections from which DA has been detected. Sinking of the spring Pseudo-nitzschia bloom may constitute a potentially important link between DA production in surface waters and benthic communities in the coastal ocean near Los Angeles. Our study indicates that toxic blooms of Pseudo-nitzschia are a recurring phenomenon along one of the most densely populated coastal stretches of the SCB and that the severity and magnitude of these events can be comparable to or greater than these events in other geographical regions affected by domoic acid.     

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.     

Reproduction of Pelagia noctiluca in the central and northern Adriatic Sea

The sequence of vitellogenesis as related to size increase of the oocytes of Pelagia noctiluca (Forskål) (Scyphozoa, Semaeostomeae) was examined to assess the influence of climatic factors on its reproductive period in the central and northern Adriatic Sea. From 1981 to 1985, 5 specimens with similar diameter were sampled each month. One ovary was excised from each individual and examined both with a videoanalyzer, to count the oocytes and to check their size distribution, and through histology to assess the stage of maturation in relation to the diameter. Oocytes of all stages were present in the ovaries of individuals down to a bell diameter of 3.5 cm. Thus reproduction occurs throughout the year in the central and north Adriatic Sea. However, the number of oocytes in different developmental stages in a gonad may range from a minimum value recorded in summer to two peaks in spring and autumn. The quantitative distribution is related to sea temperature and thus to metabolic rate, and to food availability.     

Consumption rates and prey preference of the invasive gastropod Rapana venosa in the Northern Adriatic Sea

The alien Asian gastropod Rapana venosa (Valenciennes 1846) was first recorded in 1973 along the Italian coast of the Northern Adriatic Sea. Recently, this predator of bivalves has been spreading all around the world oceans, probably helped by ship traffic and aquaculture trade. A caging experiment in natural environment was performed during the summer of 2002 in Cesenatico (Emilia-Romagna, Italy) in order to estimate consumption rates and prey preference of R. venosa. The prey items chosen were the Mediterranean mussel Mytilus galloprovincialis (Lamarck 1819), the introduced carpet clam Tapes philippinarum (Adams and Reeve 1850), both supporting the local fisheries, and the Indo-Pacific invasive clam Anadara (Scapharca) inaequivalvis (Bruguière 1789). Results showed an average consumption of about 1 bivalve prey per day (or 1.2 g wet weight per day). Predation was species and size selective towards small specimens of A. inaequivalvis; consumption of the two commercial species was lower. These results might reduce the concern about the economical impact on the local bivalve fishery due to the presence of the predatory gastropod. On the other hand, selective predation might probably alter local community structure, influencing competition amongst filter feeder/suspension feeder bivalve species and causing long-term ecological impact. The large availability of food resource and the habitat characteristics of the Emilia-Romagna littoral makes this area an important breeding ground for R. venosa in the Mediterranean Sea, thus worthy of consideration in order to understand the bioinvasion ecology of this species and to control its likely further dispersal.     

Morphology and taxonomy of chain-forming species of the genus Cochlodinium (Dinophyceae)

The morphology of an unarmored chain-forming harmful dinoflagellate Cochlodinium polykrikoides and its similar species such as Cochlodinium catenatum, Cochlodinium fulvescens, and Cochlodinium convolutum was carefully observed, emphasizing the single cell stage for clarifying taxonomically important morphological features. To differentiate C. polykrikoides from C. convolutum, the shape and the position of the nucleus are useful characters. C. polykrikoides also differs from C. fulvescens in being smaller in size, possessing many rod-shaped chloroplasts and having the sulcus running just below the cingulum on the dorsal surface. Careful observation of the ichnotype of C. catenatum suggests that C. catenatum sensu Kofoid and Swezy collected from off La Jolla, CA, USA, is not identical to C. catenatum sensu Okamura and is probably a different species, in having no chloroplasts and a nucleus positioned at the center of the cell. In addition, C. polykrikoides has many morphological features in common with C. catenatum sensu Okamura except for slightly elongate cells and is probably a junior synonym of this species.     

Relationships between occurrences of toxic Dinophysis species (Dinophyceae) and small phytoplanktons in Japanese coastal waters

Fluctuations of the genus Dinophysis, which contained several toxic species of diarrhetic shellfish poisoning (DSP), were investigated during blooms in Hiroshima Bay, Mutsu Bay and Ise Bay, Japan. The co-occurrences of small phytoplanktons (cryptophytes, other nanophytoplanktons, cyanobacteria and eukaryotic picophytoplanktons) were investigated to search for relationships with mixotrophic Dinophysis. Cryptophytes were divided into three size-groups based on length of their chloroplasts (>10, 5–10 and <5 μm) during counting by epifluorescence microscopy. Clear relationships were not found between the occurrences of Dinophysis spp. and nanophytoplanktons, cyanobacteria and eukaryotic picophytoplanktons. However, the fluctuations of small-sized cryptophytes (<5 μm) showed a close relationship with that of D. acuminata in Hiroshima Bay. In Mutsu Bay, small-sized cryptophytes also accompanied the first occurrence peak of Dinophysis spp. In Ise Bay, peaks of the occurrences of middle- and small-sized cryptophytes were observed 2–3 weeks before the peak of D. acuminata. These cryptophytes decreased rapidly with increase in D. acuminata. These results suggest the possibility that small-sized cryptophytes may be food organisms for mixotrophic Dinophysis, with the abundance of Dinophysis dependent on these cryptophytes.     

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.     

Short-term hypoxia modulates Rapana venosa (Muricidae) prey preference in Adriatic lagoons

We studied the effect of short-term hypoxia on predation by the non-native rapa whelk (Rapana venosa) on the three most common bivalve prey in northern Adriatic lagoons: the non-native ark shell (Scapharca inaequivalvis) and Manila clam (Tapes philippinarum), and the native cockle Cerastoderma glaucum. In mesocosm experiments, bivalves were offered to predator as single prey, and as multiple choice of prey. Bivalve prey showed behavioral responses when exposed to short-term hypoxia. Under normoxia, prey were consumed in almost equal numbers when offered separately, but when choice was offered to the predator R. venosa showed a marked preference for S. inaequivalvis. Short-term hypoxia increased the vulnerability of T. philippinarum, and, consequently, rapa whelks modified their predatory behavior and switched their preference to T. philippinarum. We hypothesize this to be a case of environmentally-mediated, non-reciprocal indirect interaction between the two non-native prey species, whose causes are based upon differences in specific traits of prey. Hypoxia may facilitate the coexistence of the two non-native bivalves via predator switching. The native cockle seems the net loser of the game.     

Spawning of the colonial coral <Emphasis Type="Italic">Cladocora caespitosa</Emphasis> (Anthozoa,Scleractinia) in the Southern Adriatic Sea

Data on sexual reproduction of scleractinian coral species living in temperate zones, particularly in the Mediterranean Sea, are quite scarce. This paper describes sexual reproduction of the colonial coral Cladocora caespitosa from Veliko jezero (Mljet Island) in the Adriatic Sea. Spawned orange eggs and white sperm bundles were observed on the coral bank of C. caespitosa two nights before the full moon (20 June 2005) coinciding with increasing water temperature and correlated with the lunar cycle. Spawning was observed during five nights, involving about 30% of the colonies from the coral bank. Different colonies on the bank released only one type of gamete during the reproductive period. The diameter of the sperm bundles ranged from 100 to 200 μm (average 163 μm; SD = 47.08), while the female gametes diameter ranged from 300 to 500 μm (average 416 μm; SD = 73.12).