Effect of seasonal hypoxia on the benthic foraminiferal community of the Louisiana inner continental shelf: The 20th century record

A species census in sediment core samples reveals significant changes in the composition of the Louisiana-shelf benthic foraminiferal community in the past century; these changes can be explained by an increase in the severity of seasonal hypoxia in bottom waters. Agglutinated and porcelaneous orders living in water depths less than 60 m suffered a noticeable decline during this time. In particular, the genus Quinqueloculina was severely affected by the progression of hypoxia, and nearly disappeared from parts of the study area. In contrast, several hyaline taxa, especially Nonionella basiloba, Buliminella morgani, and Epistominella vitrea, tolerated the progressive oxygen depletion well. Results of cluster and principal components analyses of the foraminiferal assemblage data match the observed species distribution trends and indicate that seasonal hypoxia on the Louisiana shelf, related to eutrophication and water stratification, worsened in the past century, even near the outer edge of the present-day zone of spring and summer oxygen depletion. The temporal trends in the foraminiferal record correspond to that of fertilizer use in the U.S. and nitrogen loading in the Mississippi River, suggesting that the anthropogenic factor has been particularly strong in the development of coastal hypoxia since the early 1940s.     

Recent benthic foraminiferal assemblages in deep high-energy environments from the Gulf of Cadiz (Spain)

The benthic environment in the Gulf of Cadiz, north-eastern Atlantic, is strongly affected by the Mediterranean outflow water undercurrent (MOW) which flows northwards along the western Iberian Margin at 500–1500 m water depth. Foraminiferal census counts of living and dead assemblages from 27 surface samples ranging from 103 to 1917 m water depth, and the examination of hard substrates reveal a close correlation of the fauna with the local hydrography and sediment facies. Four different faunal groups are separated by factor analysis of the living fauna. Assemblage 1 contains typical lower slope species and dominates samples from the lower MOW core layer and in the North Atlantic deep water below. Shelf edge foraminifera are common in assemblage 2a which shows the highest proportions in samples from 103 to 272 m. Assemblage 2b is dominated by upper slope species and suspension-feeders that are frequent in the upper MOW core layer and in distal settings between 396 and 901 m. Species from assemblage 3 prefer epibenthic habitats and are recorded with high proportions exclusively in the immediate flow paths of the upper MOW between 496 and 881 m. Colonisation structures and species composition of epibenthic assemblages from the proximal facies largely differ from those in distal settings. In general, epibenthic foraminifers only use elevated substrates under the influence of near-bottom flow. Under high current velocities, epibenthic foraminifers prefer large and heavy objects. They colonise high attachment levels where a maximum yield of advected food particles can be achieved. In distal settings at lower flow velocities, the elevation height does not exceed 20 mm above the surrounding sediment surface. This level is related to a hydrologic transition layer with high concentrations of suspended particles. The comparison of microhabitat preferences and faunal structure under high and low current velocities reveal that substrate stability may be a confining environmental variable for endobenthic and shallow epibenthic foraminifers. The observations also indicate that the preferential settling height of epibenthic foraminifera is related to the highest lateral flux rates of food particles within reach from the sea floor. A dynamic selection of elevated microhabitats is only used by 7.8% of all species recognised in the Gulf of Cadiz area.     

Distribution of the twenty seven dominant species of shelf benthic foraminifers on the continental shelf, north of Dakar (Senegal)

One hundred and thirty-one samples were collected on the shelf and on the uppermost slope (down to 180 m) off Northern Senegal, between 14 °50′ and 16 °20′ N latitude.Two hundred and three species of benthic foraminifers were recognized. Most of them were reported in previous works, along the coast of Africa. Twenty seven species are selected for detailed analysis and distribution maps are given for each of them. Q mode and R mode statistical analyses were performed on these selected species. The distribution is mainly related to depth; the second factor of distribution is the nature of the sediments: about 30% of the study area is covered by the muddy sediments of the pro-delta of the Senegal river. Hierarchical analysis shows that the main limit in the pattern of change of these 27 species with depth is about 60–65 m.A depth index Di, calculated on the basis of twelve selected species, gives a depth indication with a precision of about 40 m over the shelf. It is possible to envisage a possible use of this index for the investigation of Quaternary sea-level changes.     

Estimating denitrification in North Atlantic continental shelf sediments

A model of coupled nitrification/denitrification was developed for continental shelf sediments to estimate the spatial distribution of denitrification throughout shelf regions in the North Atlantic basin. Using data from a wide range of continental shelf regions, we found a linear relationship between denitrification and sediment oxygen uptake. This relationship was applied to specific continental shelf regions by combining it with a second regression relating sediment oxygen uptake to primary production in the overlying water. The combined equation was: denitrification (mmol N m^–2 d^–1)=0.019^* phytoplankton production (mmol C m^–2 d^–1). This relationship suggests that approximately 13% of the N incorporated into phytoplankton in shelf waters is eventually denitrified in the sediments via coupled nitrification/denitrification, assuming a C:N ratio of 6.625:1 for phytoplankton. The model calculated denitrification rates compare favorably with rates reported for several shelf regions in the North Atlantic.The model-predicted average denitrification rate for continental shelf sediments in the North Atlantic Basin is 0.69 mmol N m^{– 2} d^–1. Denitrification rates (per unit area) predicted by the model are highest for the continental shelf region in the western North Atlantic between Cape Hatteras and South Florida and lowest for Hudson Bay, the Baffin Island region, and Greenland. Within latitudinal belts, average denitrification rates were lowest in the high latitudes, intermediate in the tropics and highest in the mid-latitudes. Although denitrification rates per unit area are lowest in the high latitudes, the total N removal by denitrification (53 × 10¹⁰ mol N y^–1) is similar to that in the mid-latitudes (60 × 10¹⁰ mol N y^–1) due to the large area of continental shelf in the high latitudes. The Gulf of St. Lawrence/Grand Banks area and the North Sea are responsible for seventy-five percent of the denitrification in the high latitude region. N removal by denitrification in the western North Atlantic (96 × 10¹⁰ mol N y^–1) is two times greater than in the eastern North Atlantic (47 × 10¹⁰ mol N y^–1). This is primarily due to differences in the area of continental shelf in the two regions, as the average denitrification rate per unit area is similar in the western and eastern North Atlantic.We calculate that a total of 143 × 10¹⁰ mol N y^–1 is removed via coupled nitrification/denitrification on the North Atlantic continental shelf. This estimate is expected to underestimate total sediment denitrification because it does not include direct denitrification of nitrate from the overlying water. The rate of coupled nitrification/denitrification calculated is greater than the nitrogen inputs from atmospheric deposition and river sources combined, and suggests that onwelling of nutrient rich slope water is a major source of N for denitrification in shelf regions. For the two regions where N inputs to a shelf region from onwelling have been measured, onwelling appears to be able to balance the denitrification loss.     

东海大陆架鱼类群落的空间结构

根据1997～2000年东海大陆架水深30～200米海域4个航次的底拖网调查资料,运用多元分析(聚类分析和非度量多维标度(NMDS))、SIMPER以及BIO-ENV方法分析了东海鱼类群落的空间格局以及与非生物环境因子之间的关系。结果表明,东海大陆架海域鱼类群落在空间上可分为3个群落类型,分别为东海近海群落、东海大陆架外缘群落以及这两个群落之间的东海大陆架混合群落。东海大陆架鱼类群落的种类组成沿着水深梯度的变化明显,每种鱼类均有一定的水深范围。不同的流系具有不同的温盐特征,影响了鱼类的分布及其洄游,从而控制了不同鱼类群落类型的种类组成。东海大陆架鱼类群落类型中,大陆架外缘群落类型各调查季节在空间分布和种类组成上相对稳定,而东海近海群落类型和东海大陆架混合群落类型在夏季融合在一起,其它季节保持各自的空间结构。同时东海陆架区只在一种群落类型中出现的鱼类相对较少,多数鱼类同时出现在两个群落类型中或3个群落类型中,但是它们相对较集中的区域明显不同,在不同群落类型中的生物量和出现频率有很大的差异,这些鱼类在其不同的生长发育阶段由于对海洋环境的不同需求,它们根据繁殖或摄食需求而在东海陆架区范围内洄游以寻找最适的海洋环境,因此对3个群落类型定义是相对的,很难严格确定一个永久的鱼类群落地理区域,以形成相互隔离的鱼类群落,而且鱼类群落之间的边界随着季节不同发生变化。     

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.     

南海西北部陆架区鱼类的种类组成与群落格局

根据2006-2007年在南海西北部陆架区海域进行的4航次底拖网调查数据,对该海域鱼类的种类组成和群落格局进行分析.调查共采获鱼类262种,隶属于20目102科.以冬季出现种类数最多,为166种;夏季和秋季次之,分别为161种和155种;春季最少,为135种.运用聚类分析和非度量多维标度(NMDS)方法分析了南海西北部陆架区鱼类群落结构的空间分布.研究表明,该海域鱼类可划分为5个群落,分别为粤西沿岸群落(群落Ⅰ)、海南岛东南沿岸群落(群落Ⅱ)、海南岛东部深海群落(群落Ⅲ)、过渡群落(群落Ⅳ)和混合群落(群落Ⅴ).ANOSIM检验表明,各群落间鱼类组成的差异极为显著(R=0.847-0.939,P＜0.001).鱼类群落格局的变化与海流和水深关系密切,有较明显的沿水深梯度分布的规律.咸淡水鱼类在雷州半岛东部的冷涡区域,形成群落Ⅰ;南海暖流控制区域内的深海鱼类,形成群落Ⅲ;在南海暖流最为强劲的冬季,研究区域内大部分鱼类集结为混合性群落Ⅴ;琼东南沿岸的上升流区内形成群落Ⅱ;沿岸流强劲的夏、秋季,在沿岸流和上升流交汇处,形成过渡性群落Ⅳ.     

The Mechanism Controlling Plant Nutrient Concentrations in the Northern Adriatic Sea

A 20 year data set for the northern Adriatic was analyzed and the factors establishing the nutrient environment identified. Concentrations ranged widely (TIN 0.0–78, PO₂ 0.01–1.1, and SiO₄ 0.0–59 mmol m⁻³). In early winter remineralization increased concentrations. Characteristic winter, late spring and fall phytoplankton blooms alternately decreased and increased concentrations, as modified by river input. In summer nutrients were minimal under a semi-closed circulation pattern and high vertical stability, due to closely coupled nitrogen and phosphorus assimilation-regeneration processes and biogenic silica sedimentation. “New” primary production supported mainly by river input of “new” nutrients approximated “regenerated” primary production supported by regenerated nutrients, making the ecosystem especially sensitive to eutrophication pressure from anthropogenic increases in the Po River nutrient load.     

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.     

Live (Rose Bengal stained) and dead benthic foraminifera from the oxygen minimum zone of the Pakistan continental margin (Arabian Sea)

Live (Rose Bengal stained) and dead benthic foraminiferal communities (hard-shelled species only) from the Pakistan continental margin oxygen minimum zone (OMZ) have been studied in order to determine the relation between faunal composition and the oxygenation of bottom waters. Samples were taken from 136 m to 1870 m water depth during the intermonsoon season of 2003 (March–April). Live foraminiferal densities show a clear maximum in the first half centimetre of the sediment only few specimens are found down to 4 cm depth. The faunas exhibit a clear zonation across the Pakistan margin OMZ. Down to 500 m water depth, Uvigerina ex gr. U. semiornata and Bolivina aff. B. dilatata dominate the assemblages. These taxa are largely restricted to the upper cm of the sediment. They are adapted to the very low bottom-water oxygen values (≈ 0.1 ml/l in the OMZ core) and the extremely high input of organic carbon on the upper continental slope. The lower part of the OMZ is characterised by cosmopolitan faunas, containing also some taxa that in other areas have been described in deep infaunal microhabitats. The contrast between faunas typical for the upper part of the OMZ, and cosmopolitan faunas in the lower part of the OMZ, may be explained by a difference in the stability of dysoxic conditions over geological time periods. The core of the OMZ has been characterised by prolonged periods of stable, strongly dysoxic conditions. The lower part of the OMZ, on the contrary, has been much more variable over time-scales of 1000s and 10,000 years because of changes in surface productivity and a fluctuating intensity of NADW circulation. We suggest that, as a consequence, well-adapted, shallow infaunal taxa occupy the upper part of the OMZ, whereas in the lower part of the OMZ, cosmopolitan deep infaunal taxa have repeatedly colonised these more intermittent low oxygen environments.     

The effect of bioturbation by polychaetes (Opheliidae) on benthic foraminiferal assemblages and test preservation

Biological activity such as burrowing can alter benthic foraminiferal shell preservation and may also modify benthic foraminiferal assemblages by vertical mixing, inducing sediment homogenization. Here, we analyse benthic foraminiferal assemblages and taphonomy of upper Miocene marine deposits from Conil de la Frontera (Cádiz, south‐western Spain). The deposits consist of marls displaying a pervasive alternation of intensively bioturbated beds dominated by Macaronichnus segregatis traces (ichnofabric index 4–5) and non‐bioturbated beds. Benthic foraminiferal assemblages are dominated by Cibicidoides mundulus and Cibicides refulgens, indicating that the marls were deposited on an oligotrophic, well‐oxygenated upper slope. The impact of burrowing on the preservation of benthic foraminiferal tests was tested using Q‐mode cluster analysis, which found two well‐differentiated groups of samples, one including the non‐bioturbated beds and the other encompassing the bioturbated ones. Fragmentation and recrystallization account for the differentiation of these groups, both being higher in the bioturbated sediments. Aggressive chemical digestion by the Macaronichnus trace‐makers, assumed to be a polychaete worm of the family Opheliidae, etched the microfossil shells, making them more vulnerable to fragmentation. Intense bioturbation favoured the circulation of pore fluids, encouraging recrystallization. Pervasive burrowing resulted in significant vertical reworking of microfossils. As a consequence, benthic foraminiferal assemblages in the bioturbated beds were homogenized in the mixed layer; that is, the uppermost layer of the substrate totally burrowed. The alternation of bioturbated and non‐bioturbated beds reflects episodic transfer of food particles down slope from shallower parts of the shelf as well as from the continent due to storms under otherwise homogeneous oligotrophic marine conditions.     

Late Quaternary paleoenvironmental reconstruction of the Tremiti structural high (Central Adriatic Sea) from benthic foraminiferal assemblages

Foraminiferal analyses carried out on the VTR01/8 core (Tremiti Islands, Adriatic Sea) lead to the reconstruction of the paleoenvironmental evolution of the area. Although the core site is located at about 1000 km from the Po delta, this study shows that the installation of the recent sedimentological and run-off system (with a shore parallel clay-belt) is similar to that described at sites much closer to the Po delta. The statistical analysis singled out four associations that correspond to four distinct environments. The ecological preferences of the most abundant taxa that characterize each environment reveal that from the bottom of the core to − 50 cm, there is a slight increase of the water depth after which the water depth increases until the present level. Association A (that comprises subclusters A1, A2 and A3) is characterized by shallow-water taxa (e.g., Ammonia beccarii, Quinqueloculina spp. and Elphidium granosum) showing that in the past an infralittoral environment typified the Tremiti High. The subclusters of Association A show that during the deposition of the lower part of the core there had been an alternation of phases characterized by different degrees of riverine influence: the site was at first under the influence of river outflow (A2) that later bypassed the core site (A3) and then came back, although located further away than before (A1). With the rising of the sea level, the Tremiti High became only marginally affected by river influence because the core site is located at a considerable distance from the Po outlets and seaward with respect to the center of the modern mud-belt. As a matter of fact, association B (Bulimina marginata, Cassidulina carinata and Textularia spp.), which starts at − 46 cm, shows the typical characteristics of the modern mud-belt outer-shelf assemblage, although the presence of opportunistic species is limited by the distant position with respect to the Po delta.     

Nutrient (N,P, Si) fluxes between marine sediments and water column in coastal and open Adriatic

Nutrient benthic fluxes, as well as sediment phosphorus concentration at the open sea and coastal water stations of the Central and South Adriatic were studied during 1997–98. The fluxes were in the ranges: 0.16–2.67 mmol m^–2 d^–1 (silicate); –0.031–0.164 mmol m^–2 d^–1 (phosphate); –0.51–2.03 mmol m^–2 d^–1 (ammonia); and –1.32–1.62 mmol m^–2 d^–1 (nitrate + nitrite). Silicate flux showed a gradient from the coastal area to the open sea. Ammonia was the main nitrogen species in the flux at the estuary and bay stations, while the sum of nitrate and nitrite was predominant at the open sea stations. Relationships between phosphate and ammonia fluxes (r = 0.699, p<0.01) as well as phosphate and silicate (r = 0.529, p<0.01) were established.     

Testing for ecophenotypic variation in a benthic foraminifer

A method for identifying ecophenotypic variants is tested, using two morphological variants of the benthic foraminifer Reophax dentaliniformis Brady from the Baltic Sea. Results show that the example does not correspond to the model for ecophenotypic variation. This is interpreted as showing that the two forms are genetically differentiated and that the gene flow between them is restricted by the water circulation pattern in the Baltic.     

Laboratory observations of benthic foraminiferal cysts

Long-term observations on living benthic foraminifera in the laboratory were performed to investigate their behaviour and life style. We noticed that the formation of cysts or sedimentary envelopes is a common feature within many groups of foraminifera (organic walled, agglutinated and calcareous) in the laboratory. Several kinds of cyst were observed. In most cases, the entire foraminiferal shell was canopied with detritus or particles, but some specimens covered only parts of their body, such as the aperture region or pseudopods. Cysts were found attached to the glass walls of culture vessels or free in and on the sediment. Foraminifera stayed within cysts for hours to weeks. After leaving sedimentary envelopes, some specimens immediately started to build new ones, others not. The function of cyst formation observed in the present study was not clear, except one case where reproduction took place. Some monitored structures seemed not to be sedimentary cysts but unilocular agglutinated foraminifera with probably allogromiid-like organization. The formation of sedimentary envelopes seems to be a very basic character of foraminifera, suggesting a mechanism for the evolution of shells.     

Microzooplankton in the open waters of the northern Adriatic Sea from 1990 to 1993: the importance of copepod nauplii densities

Zooplankton was sampled during 39 cruises, from 1990 to 1993, at four fixed stations in the open northern Adriatic. Hydrographic factors were important in determining the abundance of the smallest and largest components of the northern Adriatic food chain during this period. Nauplii—especially those of the smallest size fractions—were the major mediators of material transfer between primary producers and higher trophic levels. There was a significant difference in the vertical distribution of nauplii size fractions between the eastern and western parts of the northern Adriatic, but not in their population density. According to multiple correlation analyses, the abundance of naupliar size fractions in the western area correlated strongly with temperature and with certain biological factors. This study confirms the important influence of the Po River and of mid-Adriatic waters on the planktonic ecosystem of the northern Adriatic. Electronic Publication     

The genus Dinophysis in the Southern Adriatic Sea

Abstract

Some preliminary results of a survey carried out along the Southern Adriatic coasts (Brindisi-S.M. Leuca) are reported in order to contribute to the knowledge of the phytoplankon community of this area. Ten Dinophysis species had been identified, four of them are known to be involved in DSP mussel contamination. Spatio-temporal distribution of Dinophysis spp. is shown.     

黄东海陆架区沉积物中磷的形态分布及生物可利用性

采用1992年Ruttenberg连续提取法(SEDEX)将黄东海陆架区沉积物中的磷分为交换态磷(Ex-P),Fe结合态磷(Fe-P),自生磷(Au-P),碎屑磷(De-P),有机磷(Or-P),分析了各形态磷的平面和垂直分布特征;利用沉积物年代序列测定的结果,探讨了柱状沉积物中不同形态磷的含量变化,并进一步分析了该区域磷形态的生物可利用性.结果表明,黄东海陆架区表层沉积物各形态磷平均含量为:Au-P(140.72 μg/g)＞De-P(59.23 μg/g)＞Or-P(32.69 μg/g)＞Fe-P(29.91 μg/g)＞Ex-P(5.92 μg/g);各形态磷在沉积时间序列上分布不同,反映了不同时期人类活动和气候环境等因子对磷埋藏量影响的不同,其中Au-P在长江口H1-18站位含量比南黄海中部3个站位要低得多;调查区表层潜在生物有效磷为13.55％左右,仅仅占沉积磷中的一小部分.     

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.     

Enzymatic polymorphisms in the Argentinian hake, Merluccius hubbsi Marini, of the Argentinian continental shelf

A survey described genetically encoded enzyme variation in 197 adult Argentinian hake, Merhccius hubhsi , collected from 10 sample sites on the Argentinian continental shelf. Four of 11 muscle enzyme loci were polymorphic ( PCM *, G3PDH *, EST-1 * and EST-2 *). Genotypes showed no significant difference between males and females. Only one of 37 tests of departure from Hardy-Weinberg equilibrium showed a significant difference. The allele frequency data were tested for genetic heterogeneity among locations. The overall significance of the total samples and among those from Patagonia Central was primarily due to heterogeneity at the EST-1 * locus. The implications of these findings in a population structure context are discussed.