The Fauna of Floating Cyanobacterial Mats in the Oligohaline Eulittoral Zone off Hiddensee (South-west Coast of the Baltic Sea)

Abstract. The fauna — including macrofauna, meiofauna, and large ciliates — of floating cyanobacterial mats in a brackish shallow-water area was studied by analysing six 20 cm² pieces of mat. Although these microbial aggregations were scarcely 1 cm thick, their total meiofauna abundance was about five times as high as in the uppermost 4 cm of the adjacent sediment. The mat fauna was dominated by harpacticoids, although large ciliates, rotifers, nematodes, and oligochaetes were also markedly more abundant than in the sediment. All species occurring in the mats were members of the surrounding sediment fauna. Out of the 47 species found, only a few, among them predominantly the harpacticoid, Mesochra lilljeborgi B oeck , 1864, and the nematode Daptonema setosum ( bütschli , 1874), accounted for the majority of the individual abundance in the mats. Both are regularly found in sulphidic biotopes near Hiddensee. As SEM micrographs revealed, the oligochaete Paranais litoralis (O. F. M üller , 1788) and the harpacticoid Cletocamptus confluens (Schmeil, 1894) were apparently optimal substrates for dense lawns of cyanobacteria. This indicates possible close interactions between the meiofauna and these microbiota. The frequent predation of histophagous ciliates on nematodes and harpacticoids, which were probably weakened by oxygen deficiency and/or high sulphide concentrations, were not only a sign of a generally neglected pathway in the food chain, but also impressively emphasized the huge variety of interactions taking place between meio- and microfauna within this specific benthic microcosm. Because of their floating character, the mats can play an important role in the dispersal of benthic fauna.     

虾池小型底栖动物的数量研究

提出１９８７年５月至１０月在黄岛养虾池对虾生长周期内对小型底栖动物数量逐月调查。指出：小型底栖动物总平均密度为１０９２．４ｉｎｄｓ·１０ｃｍ－２，自由生活海洋线虫和底栖桡足类是主要类群，其总平均密度分别为４６６．９和４５７．９ｉｎｄｓ·１０ｃｍ－２。８９．７％的小型底栖动物分布在沉积物表层２ｃｍ内。结果指明小型底栖动物被对虾捕食，并提出了捕食模式。     

Meiofauna and chlorophyll associated with Beggiatoa mats of a natural submarine petroleum seep

Previous studies at the Isla Vista oil seep have suggested that meiofauna, particularly nematodes, might be an important factor in explaining macrofaunal enrichment by making bacterial biomass available to the benthic food web. To explore this possibility, we analyzed meiofaunal abundance and microalgal pigments inside and just outside of bacterial mats at this natural oil seep.The bacterial mats occur where crude oil and natural gas are actively seeping out of the sediment; cores from within the mats contained a great deal of crude oil (up to 50 %). Meiofaunal abundances were the same in and out of the bacterial mats (averaging 1·-9 × 10⁶ individuals m^-2). However, dramatic changes in community structure were noticed. Harpacticoids made up 19 % of the fauna outside the mats but only 1 % inside. Pigment concentrations were also the same in both sites with phaeophytin dominating chlorophyll (120 compared to 29.8 mg m^-2). The variance of both microalgal pigments and meiofauna was much greater inside than outside, suggesting that the bacterial mats are a more heterogeneous environment.Although the effect of crude oil toxicity is not clear, the high abundances of microbial and meiofaunal biomass support the hypothesis of benthic enrichment via microbes and meiofauna.     

Size partitioning of microbial and meiobenthic biomass and respiration on Brown's Bank, south-west Nova Scotia

Biomass and respiration (oxygen consumption) of bacteria, microfauna, and meiofauna were measured in coarse sand sediment from Brown's Bank (172 m) off Nova Scotia, Canada. Community biomass, excluding macrofauna, had a median value of 35 mg C m⁻², dominated by bacteria (51%), microfauna (25%), and a minor meiofauna component (2·5%). Protozoan microfauna were mostly microflagellates (colourless cryptomonads). The experimental design allowed partitioning of benthic metabolism without using subtraction from whole community rates. Addition-removal experiments with fauna separated into size categories were used to construct a respiration-biomass regression for all taxa. Respiration rates for faunal groups were then calculated from their biomass in the natural sediment. Total microbial and meiofaunal community respiration had a median rate of 0·55 ml O₂ m⁻² h⁻¹ which was partitioned into median proportions of bacteria (50%) microflagellates (27%), and metazoan meiofauna (4%). Correlations among faunal biomass values from incubated vials of sediment suggested that bacteria were important prey for protozoans. With added biomass of meiofauna, protozoans also became a potentially important source of prey. The results demonstrated the significance of microflagellate protozoans in these sediments and their metabolic and trophic importance relative to meiofauna and even bacteria.     

Ecology of deep-sea meiobenthos in the western Pacific

I am deeply honored to have been awarded the 1988 Okada Prize of the Oceanographical Society of Japan. The present paper reviews my previous works regarding the ecology of deep-sea meiobenthos in the western Pacific area. The outline cen be summarized as follows:

Temporal patterns among meiofauna and macrofauna taxa related to changes in sediment geochemistry at an abyssal NE Atlantic site

Two major size classes of the sediment community, meiofauna and macrofauna, and four classes of lipid compounds, fatty acids, alkanes, alcohols and sterols, were investigated using multicorer and USNEL boxcorer samples, collected during six cruises over a two year period (September 1996 to September–October 1998), at the Porcupine Abyssal Plain ( 48° 50′N 16° 30′W, 4850 m depth) within the framework of the MAST 3 BENGAL project. This site was known to be subject to seasonality in the input of organic matter to the seafloor. Results are given for each faunal size class in terms of taxonomic structure at the level of phylum, class or order, depending on the taxon, and for the dominant faunal components in terms of density and vertical distribution. For each lipid compound class, results are given in concentration and vertical distribution. The taxonomic structure of each size class did not change within the study period. Total meiofaunal and macrofaunal densities were particularly high, probably reflecting the high quantity and quality of organic matter inputs to the site. The dominant components of the two size classes presented different temporal patterns in their responses to changes in their environment. Populations of meiofaunal species, a foraminiferan and an opheliid polychaete, which inhabit the surface or sub-surface of sediment and feed on phytodetritus, responded with a rapid increase in abundance to a pulse of organic input in summer 1996. The macrofaunal polychaetes showed a lagged response to the same event by slowly increasing in density. Other components of the sediment community, that can live deeper in the sediment, moved down the sediment column, in response to 1) the impoverishment and bioturbation of the surface layer, and 2) the downward mixing of organic matter in the sediment by larger organisms. In this study, different temporal patterns were demonstrated for the first time in different size classes of the sediment community, and in the biological and environmental parameters that were studied simultaneously.     

胶州湾大沽河口小型底栖动物分布的季节变动

Sediment samples were collected in the intertidal zone of the Dagu River Estuary, Jiaozhou Bay, China in April,July and October 2010 and February 2011 for examining seasonal dynamics of meiofaunal distribution and their relationship with environmental variables. A total of ten meiofaunal taxa were identified, including free-living marine nematodes, benthic copepods, polychaetes, oligochaetes, bivalves, ostracods, cnidarians, turbellarians,tardigrades and other animals. Free-living marine nematodes were the most dominant group in both abundance and biomass. The abundances of marine nematodes were higher in winter and spring than those in summer and autumn. Most of the meiofauna distributed in the 0–2 cm sediment layer. The abundance of meiofauna in hightidal zone was lower than those in low-tidal and mid-tidal zones. Results of correlation analysis showed that Chlorophyll a was the most important factor to influence the seasonal dynamics of the abundance, biomass of meiofauna and abundances of nematodes and copepods. CLUSTER analysis divided the meiofaunal assemblages into three groups and BIOENV results indicated that salinity, concentration of organic matter, sediment sorting coefficient and sediment median diameter were the main environmental factors influencing the meiofaunal assemblages.     

An assessment of the spatial heterogeneity of environmental disturbance within an enclosed harbour through the analysis of meiofauna and nematode assemblages

Significant spatial heterogeneity in the abundance and composition of meiofaunal and nematode assemblages was described inside the Genoa-Voltri harbour (Genoa, Italy) in relation to variation in the main environmental variables. In harbour sediments characterized by low Eh values and high organic matter concentrations, total meiofauna abundance was lower (948 ± 919 ind 10 cm⁻²), nematode individual biomass was higher (0.17 ± 0.07 μg C), kinorhynchs and tanaids were completely absent, and the nematode assemblage was dominated by the genera Terschellingia, Sabatieria (pulchra group) and Paracomesoma. In contrast, in sediment characterized by lower levels of organic pollution, meiofaunal abundance was higher (1085 ± 737 ind 10 cm⁻²), nematode individual biomass was lower (0.11 ± 0.04 μg C), kinorhynchs and tanaids were present and the nematodes were dominated by the genera Desmodora, Daptonema, Anticoma and Halalaimus.Environmental disturbance as assessed by the analysis of meiofaunal and nematode assemblages and sediment environmental variables changed significantly over a scale hundreds of meters, but did not follow a gradient from the inner to the outer harbour. Analysis of nematode assemblages is proposed as a useful tool for the identification of environmental risk areas which may assist in the development of good planning, monitoring programmes and better management of harbour ecosystems.     

On the ecology of meiofauna in an organically polluted estuarine mudflat

The structure, distribution and seasonal changes of the benthic meiofauna in an organically polluted, tidal, brackish-water mudflat in the Ems-Dollart estuary were analysed. Towards the outfall of polluted fresh water, macrofauna disappeared, numbers of meiofauna increased but the diversity of the meiofauna decreased. In the area surrounding the outfall the numbers and biomass of nematodes and oligochaetes increased rapidly in spring and remained high until autumn (c. 13 × 10⁶ individuals m⁻²; c. 2gCm⁻²). The benthic fauna comprised small numbers of species, dominated by a few fast-growing diatom-feeding nematodes (Eudiplogaster pararmatus and Dichromadora geophila) and oligochaetes (Amphichaeta sannio and Paranais litoralis). Eudiplogaster pararmatus exhibits brood care and it tolerates low salinities. Dichromadora geophila which is oviparous, behaved similarly regarding tolerances, life cycle and feeding but this species was less successful than Eudiplogaster in colonizing the mudflats near the outfall. The success of the two naidid oligochaete species results from their method of reproduction by means of binary fission.Most organisms fed on benthic diatoms. In spite of intensive mineralization in the mudflat, only one bacteria-feeding organism was found in abundance (the nematode Leptolaimus papilliger). The absence of macrofaunal organisms, e.g. Nereis diversicolor, is probably also responsible for the development of high densities of meiofauna in the upper sediment layers of the mudflats in the vicinity of the outfall.The heavy load of organic waste entering the Dollart reduced the diversity within the benthic ecosystem, but nevertheless a simple system remained, based on the recurrent reoxidation of the sediment surface.     

Evaluation of abyssal meiobenthos in the eastern central Pacific (Clarion-Clipperton fracture zone)

Meiobenthos were sampled from 17 stations in the abyssal deep-sea system of the central Pacific centered around 14°N, 130°W at depths 4960–5154m, during the Nixo 47 R/V Jean Charcot cruise. Meiofaunal density range from 45–89 ind. 10cm². Predominant taxa are nematodes (84–100%) and copepods (0–10%). Rotifera, Polychaeta, and Acarina also occur. Nematodes are uniformly distributed spatially with 45 species or so; Monhysteridae is the dominant taxon, and Syringolaimus sp. (Ironidae) co-occurs faithfully. Low biomass (0.4–70.6μg 10cm²) are attributed to supposed dwarfism of metazoan meiofauna and very high proportion (60–80%) of juveniles and pre-adult forms. The majority of protozoans and metazoans are detritus- or deposit-feeders; in addition symbiotic associations, coprophagy and gardening activities are frequent. In such an oligotrophic environment, low food supply may limit meiofaunal abundance, biomass and maturation, and to a lesser extent species richness.     

Response of Meiofauna to Immediate Benthic Disturbance in the Central Indian Ocean Basin

Abstract

Twenty-four stations (12 each, during pre- and postdisturbance studies) 5300 to 5330 m deep between 10°01' and 10°03'S latitude and between 75°59' and 76°02'E longitude were sampled to study the effect of benthic disturbance on the distribution of meiofauna in the Central Indian Ocean. Bottom-sampling was conducted with a box corer. Total meiofauna density ranged from 35 to 45 organisms per 10 cm² of bottom area during the predisturbance period and 21 to 32 organisms per 10 cm² during the postdisturbance period in the test and reference sites. Differences between pre- and postdisturbance study results were statistically significant (P < 0.05). Nematoda was the most abundant meta-zoan group, on an average representing >55% of the meiofaunal population. The abundance of nematodes and harpacticoid copepods as well as total meiofauna showed marked decreases during postdisturbance sampling. Vertical distribution of meiofauna in the sediment cores revealed that 75% to 90% of the metazoan population was confined to the top 2-cm layer of the sediment. Recolonization experiments suggest that harpacticoid copepods may take more time for recolonization than the nematodes and are more sensitive to the physical disturbance. These findings suggest that nematodes and harpacticoid copepods can be used as indicator organisms in recolonization experiments.     

Impact of lugworms (Arenicola marina) on mobilization and transport of fine particles and organic matter in marine sediments

We investigated the impact of sediment reworking fauna and hydrodynamics on mobilization and transport of organic matter and fine particles in marine sediments. Experiments were conducted in an annular flume using lugworms (Arenicola marina) as model organisms. The impact of lugworms on sediment characteristics and particle transport was followed through time in sediments experimentally enriched with fine particles (< 63 μm) and organic matter. Parallel experiments were run at low and high water current velocity (11 and 25 cm s^− 1) to evaluate the importance of sediment erosion at the sediment–water interface. There was no impact of fauna on sediment composition and particle transport at current velocity below the sediment erosion threshold. At current velocity above the erosion threshold, sediment reworking by lugworms resulted in dramatic particle transport (12 kg dry matter m^− 2) to an adjacent particle trap within 56 days. The transported matter was enriched 6–8 times in fine particles and organic matter when compared to the initial sediment. This study suggests that sediment reworking fauna is an important controlling factor for the particle composition of marine sediments. A. marina mediated sediment reworking greatly increases the sediment volume exposed to hydrodynamic forcing at the sediment–water interface, and through sediment resuspension control the content of fine particles and organic matter in the entire reworked sediment layer (> 20 cm depth).     

Water masses and currents of deep circulation southwest of the Shatsky Rise in the western North Pacific

We conducted full-depth hydrographic observations in the southwestern region of the Northwest Pacific Basin in September 2004 and November 2005. Deep-circulation currents crossed the observation line between the East Mariana Ridge and the Shatsky Rise, carrying Lower Circumpolar Deep Water westward in the lower deep layer (θ<1.2 °C) and Upper Circumpolar Deep Water (UCDW) and North Pacific Deep Water (NPDW) eastward in the upper deep layer (1.3–2.2 °C). In the lower deep layer at depths greater than approximately 3500 m, the eastern branch current of the deep circulation was located south of the Shatsky Rise at 30°24′–30°59′N with volume transport of 3.9 Sv (1 Sv=10⁶ m³ s⁻¹) in 2004 and at 30°06′–31°15′N with 1.6 Sv in 2005. The western branch current of the deep circulation was located north of the Ogasawara Plateau at 26°27′–27°03′N with almost 2.1 Sv in 2004 and at 26°27′–26°45′N with 2.7 Sv in 2005. Integrating past and present results, volume transport southwest of the Shatsky Rise is concluded to be a little less than 4 Sv for the eastern branch current and a little more than 2 Sv for the western branch current. In the upper deep layer at depths of approximately 2000–3500 m, UCDW and NPDW, characterized by high and low dissolved oxygen, respectively, were carried eastward at the observation line by the return flow of the deep circulation composing meridional overturning circulation. UCDW was confined between the East Mariana Ridge and the Ogasawara Plateau (22°03′–25°33′N) in 2004, whereas it extended to 26°45′N north of the Ogasawara Plateau in 2005. NPDW existed over the foot and slope of the Shatsky Rise from 29°48′N in 2004 and 30°06′N in 2005 to at least 32°30′N at the top of the Shatsky Rise. Volume transport of UCDW was estimated to be 4.6 Sv in 2004, whereas that of NPDW was 1.4 Sv in 2004 and 2.6 Sv in 2005, although the values for NPDW may be slightly underestimated, because they do not include the component north of the top of the Shatsky Rise. Volume transport of UCDW and NPDW southwest of the Shatsky Rise is concluded to be approximately 5 and 3 Sv, respectively. The pathways of UCDW and NPDW are new findings and suggest a correction for the past view of the deep circulation in the Pacific Ocean.     

Temporal changes (1989–1999) in deep-sea metazoan meiofaunal assemblages on the Porcupine Abyssal Plain,NE Atlantic

Trends among major metazoan meiofaunal taxa were investigated based on 56 deployments of a multicorer at 10 time points over a period of 11 years (1989–1999) at the Porcupine Abyssal Plain Sustained Observatory site (PAP-SO: 48°50′N 16°30′W, 4850 m depth). This area is characterised by a strong seasonality in the deposition of organic matter to the seafloor and by the massive increase in the density of holothurian species since 1996, the so-called ‘Amperima event’. Total meiofaunal densities ranged from 346 to 1074 ind.×10 cm⁻² and showed a significant increase with time when time was represented by cruises, years and the ‘Amperima period’ (1996–1999) vs. the pre-Amperima period (1989–1994). This pattern was driven mainly by the nematodes, which were the dominant taxon (∼90% of total abundance). The third most abundant group, the polychaetes, also increased significantly in abundance over the time series, while the ostracods showed a significant decrease. Most other taxa, including the second-ranked group, the copepods (harpacticoids and nauplii), did not exhibit significant temporal changes in abundance. Ordination of taxon composition showed a shift from the pre-Amperima to the Amperima periods, a trend supported by the significant correlation between the x-ordinate and time. The majority (52–75%) of meiofaunal animals inhabited the top 2 cm of the 5 cm sediment cores analysed. There were significant increases in the proportion of total meiofauna, nematodes and copepods (but not polychaetes) inhabiting the 0–1 cm layer over time (represented by cruises) and between the pre-Amperima and Amperima periods in the case of copepods and polychaetes. During the intensively sampled period (1996–1997), there were indications of seasonal changes in the vertical distribution patterns of total meiofauna and nematodes within the sediment. We discuss the potential link between temporal variations in organic matter flux to the seafloor and meiofaunal populations, considering both qualitative and quantitative changes in fluxes and how they may be linked to climate variations.     

Climate variability and pelagic fisheries in northern Chile

A time series analysis of long-term climate variability in northern Chile (18°21′–24°00′S) shows anomalies associated with the El Niño events and the longer warm period observed since 1976, followed by a cooling trend since mid 1980s. The succession of pelagic fisheries, anchovy (Engraulis ringens) and sardine (Sardinops sagax), occurring in this fishing zone was analyzed taking into account the landings, the CPUE abundance index, the fishing effort, and the environmental variables. The anchovy production model is a negative linear function of fishing effort and turbulence. For sardine, the production model is a negative linear function of fishing effort and a quadratic function of the sea surface temperature.An analysis of the relationship between recruitment, adult biomass and the environment shows that the annual recruitment of anchovy increases with turbulence intensity until wind speed reaches a value of 5.46 m s⁻¹, decreasing for higher values. For sardine, the recruitment increases with turbulence intensity until 5.63 m s⁻¹, stabilizing thereafter.It is deduced that the climatic variations associated to the El Niño events affect the abundance of coastal pelagic fishes, without forgetting the most likely effects upon its distribution and the fishing effort. However, it is the long-term variability that mainly affects the fishing activity.     

Distribution of Deep-Sea Benthos in the Proposed Mining Area of Central Indian Basin

Abstract

During the first stage of the project work on the Indian Deep-Sea Environment Experiment (INDEX), the abundance and distribution of deep-sea benthos were surveyed in the Central Indian Basin for the collection of baseline data. The deep-sea community of the sediment was characterized by a moderately high standing crop and diverse fauna. The macrofaunal component was dominated by polychaetes (100% prevalence) and peracarid crustaceans, whereas the meiofauna was dominated by nematodes and harpacticoid cope-pods. The results of this study conform to the general distribution reported elsewhere. The macrofaunal abundance showed an inverse relation to the abundance of polymetallic nodules. However, the relation between meiofaunal vertical distribution and the vertical profile of the total organic matter and total labile matter was positive.     

Distribution of Deep-Sea Benthos in the Proposed Mining Area of Central Indian Basin

During the first stage of the project work on the Indian Deep-Sea Environment Experiment (INDEX), the abundance and distribution of deep-sea benthos were surveyed in the Central Indian Basin for the collection of baseline data. The deep-sea community of the sediment was characterized by a moderately high standing crop and diverse fauna. The macrofaunal component was dominated by polychaetes (100% prevalence) and peracarid crustaceans, whereas the meiofauna was dominated by nematodes and harpacticoid copepods. The results of this study conform to the general distribution reported elsewhere. The macrofaunal abundance showed an inverse relation to the abundance of polymetallic nodules. However, the relation between meiofaunal vertical distribution and the vertical profile of the total organic matter and total labile matter was positive.     

Benthic/midwater interactions on the Madeira Abyssal Plain; evidence for biological transport pathways

An investigation of biological transport processes which may link the deep ocean floor to shallow depths was carried out at a site on the Madeira Abyssal Plain, depth 5440m, position 31°17′N 25°24′W. Benthic and midwater sampling was carried out concurrently; time lapse photography provided an estimate of biological activity on the bottom.Biomass was measured throughout the water column; overall biomass was low and no evidence was found for mass upward transport resulting from biological activity. The potential for transport via individual species does exist since a number of species were distributed from the sea bed to thousands of metres above the bottom. However, theres is no direct evidence for vertical migrations through these vast depth ranges and the numbers of individuals concerned is very small. The arrival and possible predation of a large food parcel on the deep sea floor was observed.     

Meiofauna in a cold-seep community off Hatsushima,central Japan

The community structure of the bathyal meiofauna of a cold-seep community found off Hatsushima in Sagami Bay, central Japan, was compared with the community composition outside the influence of the seep, using sediments collected during dives 226 and 227 of the deep-sea submersibleShinkai 2000. The sediment from the Hatsushima seep site (HSS) was very coarse, black in color, and with an odor of hydrogen sulfide, suggesting reduced thiobiotic conditions. The sediment from the control area was well-oxygenated, fine silt. Despite the differences in the characteristics of the sediments, the abundance of meiofauna in the HSS was not very different from that in the control area. However, its composition even at the major taxonomic group level was distinct; for example, a high nematode/copepod ratio occurred in one of the samples collected at the HSS. At the species level, nematodes were less diverse at the HSS than at the control area. The composition of the nematode fauna at the HSS showed stronger affinity with that collected at the adjacent control area than with a community sampled from other deep-sea environments or another seep community in shallow water. This emphasizes that the adaptation of nematodes to the thiobiotic condition is controlled by local conditions.     

厦门凤林红树林湿地自由生活海洋线虫群落的研究

2004年冬季在集美凤林类似底质类型、盐度和潮位的海湾地段4个断面,就不同红树植物林中的小型底栖动物数量和自由生活海洋线虫群落进行了研究。结果表明:海洋线虫是凤林红树林中小型底栖动物中的绝对优势类群,占到小型底栖动物的76.1%~96.3%;从丰度来看,旧区白骨壤(Avicennia marina)林中小型底栖动物的数量较少,有污水流过的光滩数量较大;4个断面共鉴定出海洋线虫37种(分类实体单元),其中新区秋茄(Kandelia candel)林中的生物多样性指数较低,旧区白骨壤林中群落具有较高的物种多样性;从出现的物种来看,新区秋茄林、旧区白骨壤林和光滩上出现的优势种和摄食功能群的类型各不相同。从4个断面13个站位进行的聚类分析和MDS标序分析结果推断自由生活海洋线虫的群落结构与不同的红树植物形成的沉积物有关。