Antioxidant and pro-oxidant activities of the brown algae, Laminaria digitata, Himanthalia elongata, Fucus vesiculosus, Fucus serratus and Ascophyllum nodosum

The ability of Laminaria digitata, Himanthalia elongata, Fucus vesiculosus, Fucus serratus and Ascophyllum nodosum to scavenge peroxyl radicals was investigated by kinetic studies in a model system. The thermal initiated oxidation of methyl linoleate was performed at 60°C in heptanol, with or without antioxidants. When they reached 1% of the substrate, seaweed extracts exhibited antioxidant activities by extending the induction period, but they did not suppress the rate of oxygen uptake as did vitamin E. A synergistic effect occurred when both algal extracts (1.5 g L^-1) and vitamin E (0.4 mmol L^-1) were present, and the effectiveness of the combined antioxidants during the whole induction period was vitamin E effectiveness. The synergistic effect of L. digitata, however, was subject to seasonal variations: samples collected in summer were effective synergists, whereas samples collected in winter displayed a marked negative synergism. The phospholipid fractions of F. vesiculosus, F. serratus and A. nodosum, including pigments, accounted for only 6% of the total lipid fraction, and did not exhibit a large synergistic effect. The main phospholipid was not phosphatidyl ethanolamine as usually related, but phosphatidyl inositol. Fucoxanthin had some antioxidant activity per se under our experimental conditions, but did not act as a synergist of vitamin E. The most potent synergists were recognized as chlorophyll a and related compounds by the application of liquid-liquid partition and chromatography for the identification of active components. This revised version was published online in June 2006 with corrections to the Cover Date.     

Spatial variation in polyphenolic content of Ascophyllum nodosum (Fucales,Phaeophyta)

Spatial variation in polyphenolic content in annual shoots of the brown alga Ascophyllum nodosum was quantified using a hierarchical sampling design. Three sampling levels, covering distances of 10⁰–10⁶ m, were used. Comparisons were made between two areas, Tjärnö on the Swedish west-coast and the Isle of Man in the Irish Sea, with very different types of environmental conditions. No significant differences in mean polyphenolic levels were found between the two study areas (6.6% of dry mass at Tjärnö and 9.2% at the Isle of Man), whereas significant and substantial differences were found among sites within areas (range 5.7%–11.4%) and among quadrats within sites (range 3.7%–13.1 %). The extensive variation at the smaller spatial scales points out the importance of using thorough sampling procedures at all levels in large-scale studies on algal polyphenolics, e.g. biogeographical comparisons, which have been neglected in several previous studies. Moreover, the results imply that experiments on causal factors of polyphenolic variation should be designed to explain the spatial scales on which the factors are impportant. This study also investigated the relationship between polyphenolic concentration and both plant size and mean area of annual shoots. The mean area was used as an estimate of the mean growth rate of the annual shoots within an individual. No significant relationships were found between shoot growth rate, or plant size, and polyphenolic levels in annual shoots at any of the three spatial scales that were investigated.     

Expressed sequence tag-derived polymorphic SSR markers for Fucus serratus and amplification in other species of Fucus

The seaweed genus Fucus is a dominant component of intertidal shores throughout the North Atlantic and North Pacific and has been the focus of considerable developmental, ecological, and evolutionary research for the past century. Here, we present details of 21 expressed sequence tag-derived simple sequence repeat markers (microsatellites). All 21 were polymorphic for F. serratus, which also display considerable cross-reactivity with the sister species F. distichus (18) and the more distantly related F. vesiculosus (13), and F. spiralis (5).     

盐城自然保护区射阳河口潮间带大型底栖动物空间分布与季节变化

2008年11月—2009年10月,在盐城自然保护区射阳河口潮间带按月份对大型底栖动物群落进行取样。共得到大型底栖动物16种,隶属3门4纲15科,为软体动物、节肢动物甲壳类和环节动物多毛类,其中,潮间带米草丛6种,高潮带光滩7种,中潮带光滩13种,低潮带光滩7种。对潮间带各样点的物种数、密度、生物量以及不同样点的生物多样性指数Margalef指数(S)、Shannon指数(H)、Pielou指数(J)和Simpson指数(D)进行样点-季节间无重复双因素方差分析,发现样点间物种数、密度和生物量差异极显著(所有P<0.01),三者季节间差异均不显著;样点间H、D和S差异显著(所有P<0.05),季节间差异均不显著;而季节间和样点间J差异均不显著。对四季4个样点的群落进行聚类和排序分析发现,中、高潮带光滩界限不明显,潮间带米草丛和低潮带光滩界限明显。结果表明:生境条件的不同是潮间带大型底栖动物群落组成差异的主要原因,同一生境大型底栖动物群落季节间差异不显著。     

Effect of epiphytism on reproductive and vegetative lateral formation in the brown, intertidal seaweed Ascophyllum nodosum (Phaeophyceae)

In the brown alga Ascophyllum nodosum (L) Le Jolis, a common species on sheltered Northern temperate rocky shores, gametes are produced in receptacles that emerge from small depressions (lateral pits) along the branched frond. These lateral pits are also the preferred settling site for the obligate epiphyte Polysiphonia lanosa (L) Tandy. Therefore, epiphytism can be expected to interfere with host reproductive output. The present study investigated the potential impact of the epiphyte on A. nodosum in two series of laboratory experiments that measured: (i) the direct shading of the host plant underneath an epiphyte canopy; and (ii) the development of receptacles in clean and epiphytised A. nodosum segments (excised from individual fronds) over a 6 month period. These experiments showed that light reaching emerged fronds underneath a dense epiphyte cover was reduced by 40%, and this was independent of the degree of desiccation the epiphyte experienced. Concurrently, in the growth study with epiphytised A. nodosum segments (segments with one clean and one epiphytised lateral pit) total receptacle biomass per epiphytised fragment was significantly reduced compared with clean segments (0.52 g and 1.25 g per gram of frond segment, respectively), although this effect was only significant in A. nodosum from sheltered shores. However, expressed as biomass per lateral pit, receptacle biomass in the remaining clean lateral pits in epiphytised segments was significantly increased in segments from both shores, demonstrating that A. nodosum can at least partially compensate for the loss of production resulting from epiphytism.     

Diversity, biomass and distribution pattern of Sargassum beds in the South West lagoon of New Caledonia (South Pacific)

Sargassum (Phaeophyceae, Fucales) is a genus of worldwide distribution recently recognised to proliferate in several regions of the South Pacific. In New Caledonia, species of this genus naturally structure one of the major lagoon habitats but their extent, composition and biomass remain largely unknown. To fill these gaps in our knowledge over large areas, we applied a combination of remote sensing and in situ methods applied to the Neo Caledonian South West Lagoon. Space borne high resolution Landsat (30-m resolution) and Quickbird (2.4-m resolution) images of the Neo Caledonian South West lagoon were used to estimate the spatial extent of the different beds of interest. Species composition was determined for 11 Sargassum beds and seasonal variations were investigated for four representative beds over an 18 month period using quadrats and transects. Relationships between surface cover and biomass were estimated from seasonal field data sets. Finally, four methods requiring variable levels of sampling effort were designed to estimate the total biomass at the scale of each bed, considering (or not) the specific composition, and spatial and temporal variations. Seven Sargassaceae species were identified. Mean surface cover (24.4–51.6%) and total biomass [3.4–1,461.9 t dry matter (DM)] varied widely between beds. Overall, biomass temporal variations were not significantly different, but species-level variations seemed to be bed-specific. The extent of the 11 beds was 9 km²; their total biomass was estimated and compared using each of the four methods, and the most precise method provided an estimate of 2,900 t DM. This study demonstrates a way of characterising Sargassum beds, efficiently and on a large scale, using a combination of remotely sensed and in situ data. These methods should be useful for possible future biomonitoring of Sargassum beds in New Caledonia, and in other areas worldwide.     

Seasonal changes in an intertidal annual algal assemblage in northern Japan: The role of pre-emption and grazing on algal replacement

The pattern and process of seasonal changes in an intertidal annual algal assemblage were examined at Hiura, northern Japan. Short-term field experiments (<2months duration) were set up to quantify the effects of both grazing and pre-emption on species replacement in the assemblage in three different seasons. An 8-month field experiment was set up to quantify long-term effects, including the indirect effects of both grazing and competitive dominance on the community structure. Results suggested that seasonal change in the algal assemblage resulted from the interaction of abiotic environmental change, competition and grazing. The relative contribution of these factors varied within a short period, presumably as a result of seasonal changes in physical environmental stress, free space availability and grazing pressure. From February to March, when grazer density was low and there was much free space available for algae, the dominant species shifted from foliose green alga Monostroma angicava to filamentous red alga Bangia atropurpurea, because B.atropurpurea grew faster than M.angicava. This species replacement was not influenced strongly by biological interaction but by temporal changes in abiotic environmental conditions. From April to mid May, when there was less free space available for algae in the natural community, the dominant B.atropurpurea decreased with increasing foliose red alga Porphyra yezoensis, because only P.yezoensis was able to invade an area pre-empted by algae. Grazing did not affect this species replacement. After mid May, the two dominant species, P.yezoensis and B.atropurpurea, decreased. Their decline was mainly caused by desiccation stress and was partially affected by grazing.     

Spatial distribution of phytomass carbon in Indian forests

A spatial analysis of phytomass carbon (C) in Indian forests for the period (1988–94) at the district‐level is reported. District‐level forest phytomass C (1988, 1994) was computed by combining remote sensing‐based forest area inventories on 1:250 000 scale, field inventories of growing stock volume by the Forest Survey of India (FSI), and crown density‐based biomass expansion factors. The estimates of forest area inventory, forest phytomass C density, and forest phytomass C pools (1988–94) were linked to the district boundary coverage of India in ARC/INFO Geographic Information System (GIS). Of the total 386 districts examined, only 17 districts had more than 75% forest cover (as percent of their geographic area) in 1988. Estimated district‐level forest phytomass C densities ranged from 4.3 to 206.8 MgC ha^?1*. At the national level, forest phytomass C pool was estimated to be 3871.2 and 3874.3 TgC# in 1988 and 1994, respectively. Although the aggregated national estimates were similar, the district‐level change analysis indicated a decrease of 77.8 TgC and an increase of 81 TgC (1988–94). The highest decrease of 10.4 TgC was observed in Vishakapatnam district (Andhra Pradesh) while largest increase of 9.8 TgC in Bastar district (Madhya Pradesh) resulting from deforestation and afforestation activities, respectively. At the national level, the total forest cover decreased by ～0.6 Mha, however, the district‐level spatial analysis indicated an increase of 1.07 Mha, and decrease of 1.65 Mha forest cover during 1988–94 period. Although, this is the first district level phytomass C analysis of Indian forests, the aggregated results at state‐level were close to the earlier estimates. The spatial analysis identified the districts that have undergone significant changes in their forest phytomass C during the study period. This improved understanding of forest phytomass C pools is important to sustainable development and conservation of forests, mitigation strategies for C sequestration, and accurate estimation of contribution of land use changes to C emission in India.     

Predation by gulls on crabs in rocky intertidal and shallow subtidal zones of the Gulf of Maine

Most organisms in intertidal areas are marine in origin; many have distributions that extend into the subtidal zone. Terrestrial predators such as mammals and birds may exploit these animals during low tide and can have considerable effects on intertidal food webs. Several studies have shown that avian predators are capable of reducing densities of sessile and slow-moving intertidal invertebrates but very few studies have considered avian predation on mobile invertebrate predators such as crabs. In this study, we investigated predation by Great Black-backed Gulls (Larus marinus Linnaeus) on three species of crabs (Cancer borealis Stimpson, Cancer irroratus Say, and Carcinus maenas Linnaeus). The study was at Appledore Island, ME (a gull breeding island) and 8 other sites throughout the Gulf of Maine, including breeding islands and mainland sites. On Appledore Island, intertidal and subtidal zones provided over one-third of prey remains found at gull nests, and crabs were a substantial proportion (∼ 30% to 40%) of the total remains. Similarly, collections of prey remains from intertidal areas indicated that crabs were by far the most common marine prey. C. borealis was eaten far more often and C. irroratus and C. maenas less often than expected at each site. Comparing numbers of carapaces to densities of crabs in low intertidal and shallow subtidal zones at each site, we estimated that gulls remove between 15% and 64% of C. borealis during diurnal low tides. The proportion of C. borealis eaten by gulls was independent of proximity to a gull colony. Approximately 97% of the outer coast of Maine is within 20 km of a breeding island. Thus, a lot of gull predation on crabs may occur throughout the Gulf of Maine during summer months. Crabs are important predators of other invertebrates; if predation by gulls reduces the number of crabs in intertidal and shallow subtidal areas, gulls may have important indirect effects on intertidal food webs.     

Intertidal distribution and species composition of brachyuran crabs at two rocky shores in Central Portugal

The objectives of the present study are to describe and compare the brachyuran community of rocky shores within the Central Portuguese coast and to examine the zonation patterns of the most representative species. For this, randomly placed transects were surveyed to obtain crab counts according to microhabitat and intertidal level. Repeated sampling in two different shores during two different seasons provided spatial and temporal replication for zonation analyses. Seven species were registered: Pachygrapsus marmoratus, Eriphia verrucosa, Xantho incisus, Carcinus maenas, Necora puber, Pirimela denticulata and Pilumnus hirtellus. Species density rankings are the same at both localities, but the less exposed shore presents higher diversity. While most species are mainly confined to specific microhabitats in the lower level, P. marmoratus and E. verrucosa can exploit the whole intertidal range. Regardless of shore and season, E. verrucosa is more abundant in the lower intertidal levels, while no such zonation patterns were recorded for P. marmoratus. Initial predictions concerning the effect of wave exposure and temperature on the zonation of those species are not validated after analysing the factorial model proposed. Between-shore contrasts were found instead, with higher densities recorded in the more exposed locality for both species. Possible causes of the observed patterns are discussed.     

Advanced phenotyping offers opportunities for improved breeding of forage and turf species

Background and Aims

Advanced phenotyping, i.e. the application of automated, high-throughput methods to characterize plant architecture and performance, has the potential to accelerate breeding progress but is far from being routinely used in current breeding approaches. In forage and turf improvement programmes, in particular, where breeding populations and cultivars are characterized by high genetic diversity and substantial genotype × environment interactions, precise and efficient phenotyping is essential to meet future challenges imposed by climate change, growing demand and declining resources.

Scope

This review highlights recent achievements in the establishment of phenotyping tools and platforms. Some of these tools have originally been established in remote sensing, some in precision agriculture, while others are laboratory-based imaging procedures. They quantify plant colour, spectral reflection, chlorophyll-fluorescence, temperature and other properties, from which traits such as biomass, architecture, photosynthetic efficiency, stomatal aperture or stress resistance can be derived. Applications of these methods in the context of forage and turf breeding are discussed.

Conclusions

Progress in cutting-edge molecular breeding tools is beginning to be matched by progress in automated non-destructive imaging methods. Joint application of precise phenotyping machinery and molecular tools in optimized breeding schemes will improve forage and turf breeding in the near future and will thereby contribute to amended performance of managed grassland agroecosystems.     

Monitoring Karenia brevis blooms in the Gulf of Mexico using satellite ocean color imagery and other data

Harmful algal blooms (HABs) of Karenia brevis are a recurrent problem in the Gulf of Mexico, with nearly annual occurrences on the Florida southwest coast, and fewer occurrences on the northwest Florida and Texas coasts. Beginning in 1999, the National Oceanic and Atmospheric Administration has issued the Gulf of Mexico HAB Bulletins to support state monitoring and management efforts. These bulletins involve analysis of satellite imagery with field and meteorological station data. The effort involves several components or models: (a) monitoring the movement of an algal bloom that has previously been identified as a HAB (type 1 forecast); (b) detecting new blooms as HAB or non-HAB (type 2); (c) predicting the movement of an identified HAB (type 3); (d) predicting conditions favorable for a HAB to occur where blooms have not yet been observed (type 4). The types 1 and 2 involve methods of bloom detection requiring routine remote sensing, especially satellite ocean color imagery and in situ data. Prediction (types 3 and 4) builds on the monitoring capability by using interpretative and numerical modeling. Successful forecasts cover more than 1000 km of coast and require routine input of remotely sensed and in situ data.The data sources used in this effort include ocean color imagery from the Sea-Viewing Wide Field-of-View Sensor/OrbView-2 satellite and processed using coastal-specific algorithms, wind data from coastal and offshore buoys, field observations of bloom location and intensity provided by state agencies, and forecasts from the National Weather Service. The HAB Bulletins began in coordination with the state of Florida in autumn of 1999 and included K. brevis bloom monitoring (type 1), with limited advisories on transport (type 3) and the detection of blooms in new areas (type 2). In autumn 2000, we improved both the transport forecasts and detection capabilities and began prediction of conditions favorable for bloom development (type 4). The HAB Bulletins have had several successes. The state of Florida was advised of the potential for a bloom to occur at the end of September 2000 (type 4), and the state was alerted to the position of blooms in January 2000 and October 2001 in areas that had not been previously sampled (type 3). These successful communications of HAB activity allowed Florida agencies responsible for shellfish management and public health to respond to a rapidly developing event in a timely, efficient manner.     

Biomass, species composition and diversity of epipelic algae in mire pools

The biomass, species composition and diversity of epipelic algae in two small pools of contrasting physicochemical characteristics in Miyatoko Mire were studied in 1992 (pool 3 =site B4 and pool 50 =site D2). A total of 93 species and 67 species of epipelic algae occurred at sites B4 and D2, respectively. Considerable differences were observed between the two sites in the seasonal fluctuations of species number, biomass and dominant species. At site B4, little changed with species number during April–August and markedly increased in October, while biomass was largest in April and gradually decreased during June–October. Diatoms and desmids occupied 33–82% and 15–63% of total algal biomass, respectively. At site D2, species number and biomass were small in April just after the snow-thaw, and increased in June and decreased in August and October. Diatoms occupied 90–98% of total algal biomass. The species diversity was always higher at site D2 than B4. As a result of analyses of water chemistry in the two pools, pool B4 can be recognized as a habitat experiencing high disturbance frequency. It is predicted that pools experiencing frequent disturbance will have less epipelic algal biomass and diversity.     

Spatial distribution of cadmium in leaves of metal hyperaccumulating Thlaspi praecox using micro-PIXE

* Localization of cadmium (Cd) and other elements was studied in the leaves of the field-collected cadmium/zinc (Cd/Zn) hyperaccumulator Thlaspi praecox from an area polluted with heavy metals near a lead mine and smelter in Slovenia, using micro-PIXE (proton-induced X-ray emission). * The samples were prepared using cryofixation. Quantitative elemental maps and average concentrations in whole-leaf cross-sections and selected tissues were obtained. * Cd was preferentially localized in the lower epidermis (820 microg g(-1) DW), vascular bundles and upper epidermis, whereas about twice the lower concentrations were found in the mesophyll. * Taking into account the large volume of the mesophyll compared with the epidermis, the mesophyll is indicated as a relatively large pool of Cd, possibly involved in Cd detoxification/dilution at the tissue and cellular level.     

遥感技术支持下的植被生产力与生物量研究进展

目前广泛应用于植被生产力与生物量估算的遥感模型主要有经验模型、物理模型、半经验模型和综合模型 ,它们的应用受到诸如大气、背景、地形、植被覆盖率与结构等因素的影响。遥感技术的迅速发展及其它技术的应用 ,包括热红外、微波和激光遥感仪器以及多角度、高光谱和高分辨率技术等 ,正逐步消除或降低影响因素 ,进一步提高植被生产力与生物量估算的范围和精度     

Mapping species density of trees,shrubs and vines in a tropical forest,using field measurements,satellite multiespectral imagery and spatial interpolation

We estimated the number of species in a tropical forest landscape in Quintana Roo, Mexico, based on the relationship between reflectance values of satellite imagery and field measurements of plant species density (mean number of species per plot). Total species density as well as that of tree, shrub and vine species were identified from 141 sampling quadrats (16543 individuals sampled). Spatial prediction of plant diversity was performed using universal kriging. This approach considered the linear relationship between plant species density and reflectance values of Thematic Mapper™, as well as the spatial dependence of the observations. We explored the linear relationships between spectral properties of TM bands and the species density of trees, shrubs and vines, using regression analysis. We employed Akaike Information Criterion (AIC) to select a set of candidate models. Based on Akaike weights, we calculated model-averaged parameters. Linear regression between number of species and reflectance values of TM bands yielded regression residuals. We used variogram analysis to analyze the spatial structure of these residuals. Results show that accounting for spatial autocorrelation in the residual variation improved model R² from 0.57 to 0.66 for number of all species, from 0.58 to 0.65 for number of tree species, from 0.26 to 0.41 for number of shrub species and from 0.13 to 0.17 for species density of vines. The empirical models we developed can be used to predict landscape-level species density in the Yucatan Peninsula, helping to guide and evaluate management and conservation strategies.     

Spatial distribution and species composition of prosthecate bacteria in Lake Baikal

From the water column of Lake Baikal, several strains of prosthecate bacteria belonging to the genera Caulobacter and Brevundimonas were isolated. In this article, the methods applied for their isolation and cell number determination are described; the occurrence frequency and spatial distribution of these microorganisms in the lake are demonstrated. Characterization of the species composition of cultivable and uncultivable prosthecate bacteria was carried out using the methods of traditional and molecular microbiology, respectively. A comparative phylogenetic analysis of the DNA sequences of uncultivable bacteria, which showed homology to the members of the alpha subclass of proteobacteria, was carried out. It was demonstrated that the lake water column is inhabited by uncultivable alpha-proteobacteria of uncertain phylogenetic affinity, in addition to representatives of the species Caulobacter vibrioides and C. leidyi, which were detected by traditional microbiological methods.     

夜郎湖水库水体不同形态汞的时空分布

于2006年7月(夏季)、2007年1月(冬季)和3月(春季)采集了贵州省夜郎湖水库水样,研究了不同形态汞(总汞、溶解态汞、颗粒态汞)的时空分布特征及其影响因素.结果表明,夏季水体总汞、溶解态汞、颗粒态汞平均含量分别为4.48±2.59、2.37±1.40、2.11±1.86 ng·L-1,均显著高于冬季和春季(P＜0.001),而冬春2季不同形态汞含量无明显差异.水质参数悬浮颗粒物(SPM)和硝酸盐(NO-3)与不同形态汞之间均存在显著的正相关关系,表明这些参数对于不同形态汞的季节分布起着重要作用.夏季农业耕作活动相对活跃,表层土壤的扰动增加,雨水冲刷农田土壤,带进大量的外源颗粒物,致使夜郎湖水体夏季总汞水平较高.空间分布表明,夜郎湖水库夏季总汞平均浓度从水库入库河流至大坝方向、出库河流呈现总体下降的分布趋势,但水体各采样剖面没有明显的分布规律.     

Spatial predictability of juvenile fish species richness and abundance in a coral reef environment

Juvenile reef fish communities represent an essential component of coral reef ecosystems in the current focus of fish population dynamics and coral reef resilience. Juvenile fish survival depends on habitat characteristics and is, following settlement, the first determinant of the number of individuals within adult populations. The goal of this study was to provide methods for mapping juvenile fish species richness and abundance into spatial domains suitable for micro and meso-scale analysis and management decisions. Generalized Linear Models predicting juvenile fish species richness and abundance were developed according to spatial and temporal environmental variables measured from 10 m up to 10 km in the southwest lagoon of New Caledonia. The statistical model was further spatially generalized using a 1.5-m resolution, independently created, remotely sensed, habitat map. This procedure revealed that : (1) spatial factors at 10 to 100-m scale explained up to 71% of variability in juvenile species richness, (2) a small improvement (75%) was gained when a combination of environmental variables at different spatial and temporal scales was used and (3) the coupling of remotely sensed data, geographical information system tools and point-based ecological data showed that the highest species richness and abundance were predicted along a narrow margin overlapping the coral reef flat and adjacent seagrass beds. Spatially explicit models of species distribution may be relevant for the management of reef communities when strong relationships exist between faunistic and environmental variables and when models are built at appropriate scales.     

Factors influencing the species composition,distribution and abundance of benthic invertebrates in the profundal zone of a eutrophic northern lake

Factors influencing the species composition, distribution and abundance of benthic invertebrates were determined in a eutrophic subarctic lake from April 1978 to April 1979. Collections were made at five stations located at depths of 4 to 13 m. The largest populations of up to 5 × 10³ animals m^–2 were found in the deepest part of the lake. of the 24 species recorded in this area, the chironomidsProcladius denticulatus, Dicrotendipes modestus, Chironomus decorus andGlyptotendipes barbipes were most common. The strong development of benthos in the profundal zone was attributed to a consistently large supply of food and warm (4 °C) winter temperatures on bottom. Slightly smaller populations (up to 4 × 10³ animals, m^–2), composed of 19–23 species, occurred in shallower water, a reflection of lower (1.5 °C) winter temperatures. In the anoxic northern part of the lake, only 4–8 species were found in low numbers (400–1 000 animals m^–2). This was likely due to low (<5% saturation) oxygen levels in water and high organic content (18.5%) of the sediments.