The effects of predation by juvenile fish on the meiobenthic community structure in a natural pond

相似文献   

Top-down control of a meiobenthic community by two juvenile freshwater fish species

Top-down control of prey assemblages by fish predation has been clearly demonstrated for zooplankton and macroinvertebrates. However, in the benthic communities of freshwater ecosystems, the impact of fish predation on meiofaunal assemblages is nearly unknown. In this study, the predation effects of juvenile carp (Cyprinus carpio) and gudgeon (Gobio gobio) on meiofaunal abundance, biomass, community structure, and the diversity of nematodes were examined using microcosms that were sampled repeatedly over 64 days. Significant differences in abundance and biomass were found between the two fish treatments (carp and gudgeon) and their respective controls for nematodes, oligochaetes, and crustaceans (copepods, harpacticoids, ostracods, and cladocerans), but not for rotifers. These changes were consistent with top-down control of the freshwater meiofaunal assemblages in the microcosms over time. By contrast, small-bodied meiofauna was more abundant, suggesting indirect facilitation. Neither the species richness nor the diversity of the nematode community was affected by fish predation. The results indicate that predation by juvenile freshwater fish depresses the overall abundance and biomass of meiofaunal assemblages, except for rotifers, and alters the size structure of the meiofaunal community. Therefore, the meiofaunal assemblages of freshwater ecosystems may be influenced by bottom-feeding juvenile fish, e.g., carp and gudgeon, through top-down control of meiofaunal populations.     

Experimental evidence of predation by juvenile flounder, Platichthys flesus, on a shallow water meiobenthic community

In the northern Baltic Sea juvenile flounder, Platichthys flesus, occur in high abundances on shallow sandy bottoms in late summer and autumn. They feed mainly on meiofauna and the ontogenetic switch to macrofaunal sized food occurs at a larger size here than in other areas, exemplifying the high relative importance of meiofauna. Consequently, juvenile P. flesus in the Baltic feed for a longer period on meiofauna, and could thus be expected to have a stronger predation effect on the meiofaunal assemblages. In this study the predation effects of juvenile P. flesus on meiofaunal abundances and community structure were studied using microcosms that were sampled repeatedly over a 3-week period. Significant differences between treatment and control were found for the total number of taxa, for abundances of harpacticoids, copepod nauplii and ostracods. The nematode community was not affected, but one genus, Axonolaimus, was negatively affected by predation. The predation affected meiofaunal community structure as the major taxon diversity was significantly reduced. The results suggest that the meiofauna on shallow sandy bottoms may be structured by juvenile P. flesus, and the combined predation pressure of juvenile flounder and other epibenthic predators in the area might be considerable. Microcosms are effective in testing natural predation, especially on meiofaunal communities, but field experiments should be conducted to account for the physical characteristics of the area studied.     

Diversity of larger consumers enhances interference competition effects on smaller competitors

Competition between large and small species for the same food is common in a number of ecosystems including aquatic ones. How diversity of larger consumers affects the access of smaller competitors to a limiting resource is not well understood. We tested experimentally how species richness (0–3 spp.) of benthic deposit-feeding macrofauna changes meiofaunal ostracods’ incorporation of fresh organic matter from a stable-isotope-labeled cyanobacterial bloom, using fauna from the species-poor Baltic Sea. Presence of macrofauna mostly decreased meiofaunal incorporation of bloom material, depending on the macrofauna species present. As expected, the species identity of macrofauna influenced the incorporation of organic matter by meiofauna. Interestingly, our results show that, in addition, species richness of the macrofauna significantly reduced meiofauna incorporation of freshly settled nitrogen and carbon. With more than one macrofauna species, the reduction was always greater than expected from the single-species treatments. Field data from the Baltic Sea showed a negative correlation between macrofauna diversity and meiofaunal ostracod abundance, as expected from the experimental results. We argue that this is caused by interference competition, due to spatial niche differentiation between macrofauna species reducing the sediment volume in which ostracods can feed undisturbed by larger competitors. Interference from macrofauna significantly reduces organic matter incorporation by meiofauna, indicating that diversity of larger consumers is an important factor controlling the access of smaller competitors to a limiting food resource.     

The effects of nutrient enrichment on a freshwater meiofaunal assemblage

1. The effects of eutrophication on phytoplankton, zooplankton and fish in lakes are well known. By contrast, little is known about the response of the zoobenthos to nutrient enrichment, while smaller organisms, such as the meiofauna, have for the most part been neglected. 2. In a long‐term (16 months) microcosm experiment, we assessed the effects of five levels of nutrients [total phosphorus (TP), 7–250 μg L^?1; nitrate, 2–8 mg L^?1] on a freshwater meiofaunal assemblage and on nematode diversity in particular. 3. Within the first 8 months, meiofaunal succession was only weakly affected, whereas, during the last 4 months, nutrient addition influenced most of the main taxa, with a concomitant change in the assemblage structure. 4. The density of the numerically dominant nematodes decreased upon nutrient enrichment, whereas ostracods became more numerous. Other taxa, including copepods, reached a maximum at intermediate nutrient levels or, in case of oligochaetes, were almost unaffected by nutrient enrichment. However, the changes in the density of the main taxa were usually insufficient to alter their biomass. Consequently, meiofaunal biomass was remarkably unresponsive to nutrient addition, while meiofaunal density displayed a unimodal relationship, with a peak at a TP concentration of 30 μg L^?1. In addition, nematode species richness decreased significantly with increasing nutrient concentrations. 5. We hypothesise that the response of meiofaunal taxa to nutrients is attributable to the development of primary producers, which shifted with enrichment from low densities of edible diatoms and unicellular green algae to large standing stocks of inedible forms, such as Lemna minor and Cladophora spp.     

Recolonisation of meiofauna after catastrophic iceberg scouring in shallow Antarctic sediments

A series of 10 samples from sediment in and adjacent to a shallow coastal iceberg scour at Signy Island, Antarctica, were taken by hand coring from 17 December 1993 until 23 August 1994. Scouring by the iceberg led to more than a 95% decrease in meiofaunal abundance and to a certain degree of reduction in diversity. Nematodes were always the most dominant group of meiofauna. The return of major meiofauna groups to control levels was accomplished in 30 days, although a decrease in abundance on the 50th day made interpretation difficult. The pioneering meiofaunal colonisers were copepods and ostracods, followed by nematodes. Microlaimus sp.1 was dominant among the nematodes throughout the whole period. Epistratum feeders and non-selective deposit feeders were highly dominant over selective deposit feeders and predators/omnivores. The Maturity Index, a measure for stress within nematode communities, was relatively low at all times and in controls, which indicates that r-strategists prevail in this community. In spite of the catastrophic destruction, nematode community structure was not affected by the iceberg impact, and there was no evidence of succession during recovery. This suggests that the nematode community in the shallow subtidal coast at Signy Island is well adapted to ice disturbance.     

Distribution,abundance and ecology of the meiofauna in a tropical estuary along the west coast of India

The distribution and abundance of subtidal meiofauna in Mandovi estuary of Goa were studied from June 1983 to June 1984. Monthly faunal abundance ranged from 491 to 2791/10 cm² and dry weight biomass from 0.16 to 2.80 mg 10 cm². Free living nematodes were the dominant group contributing over 75% of the total density and 30 to 42% of the total biomass. Among nematodes the deposit feeders were more abundant in fine muddy substratum while epigrowth feeders dominated in sandy substratum.Harpacticoids were next, comprising 6.9 to 8.7% of the total meiofauna number, followed by turbellaria (3.8–4.5%), polychaeta (2.8–3.2%) and ostracods (1.6–4.5%) The contribution of other groups to faunal density was 4.5–6.2%. In the biomass the ostracods contributed most (29.8–54.7%), followed by nematodes (23.8–34.6%). Over 60% of the fauna occurred in the top 2 cm of the sediment and the faunal density reduced significantly with increasing depth in the sediment. The vertical distribution of meiofauna was positively correlated to the vertical distribution of Eh, chlorophyll a and interstitial water. Seasonality was greatly influenced by the south-west monsoon and the fauna quickly repopulated after the monsoon. Salinity, temperature and food influenced the faunal abundance.     

Distribution and abundance of meiofauna in intertidal sand substrata around Iceland

Iceland is situated in an important subarctic transition area where complex oceanographic dynamics occur. The intertidal, subtidal, and deep-sea faunal communities of Iceland are being intensively studied, as a critical resource for continued sustainability of fisheries and the preservation of northern littoral ecosystems. However, the meiofaunal communities and the environmental factors affecting them are still relatively poorly known. The meiobenthic metazoan community was studied with core sampling in 23 sandy beaches along the intertidal zone of the Iceland coast in a campaign developed in September 2003. Small-scale variation in meiofauna composition (major taxa) was explored and related to biotic and abiotic factors at different scales, such as beach exposure, granulometry, and organic matter content. Differences in meiofaunal community structure at a low taxonomic resolution appeared among beaches located within wide biogeographical zones of hydrobiological significance (NE and SW Shelf regions) and exposure degrees. Seventeen major taxa were recorded. In contrast with more local and taxon-focused studies, oligochaetes were the dominant group all around Iceland, followed by nematodes, turbellarians, gastrotrichs, and copepods (mainly harpacticoids). Acari, ascidians, bivalves, cnidarians, collembolans, gastropods, isopods, kinorhynchs, insects, nemerteans, ostracods, and polychaetes were relatively scarce groups, together being less than 1.6% of the meiofauna. There was a large variation in meiofaunal abundance between sites. Maximum abundances (>500 ind. cm^?2) were found in Sau?arkrökur, Hraunhafnartangi, and Skálaness, whereas minimum abundances (<40 ind. cm^?2) were recorded in Magnavík, Jokülsárlón (glacier beach site), Vikurnúpur, Breidalsvík, and Stokknes. We did not find a clear pattern in overall meiofaunal abundance regarding the degree of exposure of beaches. Oligochaetes, nematodes, and copepods were relatively more abundant in sheltered beaches, whereas turbellarians and gastrotrichs tended to be more abundant in exposed beaches. The best correlates of meiofaunal composition and abundance within beaches were the proportion of gravels and the content of utilizable organic matter in the sediment. We should consider factors operating at wider scales (importantly beach exposure and overall situation in the complex oceanographical context of Iceland) to find a pattern in the local structure of intertidal meiofaunal assemblages.     

Meiofaunal distribution in Hornsund fjord,Spitsbergen

The meiofaunal community structure at 32 stations in Hornsund fjord (77°N) was investigated, and results were compared with data from another Spitsbergen fjord, Kongsfjorden (79°N). Steep environmental gradients of sedimentation, organic matter content, and salinity from the inner to the outer basin of the fjord are present due to intensive glacial discharges of meltwater and ice. As the natural environmental disturbances were described for macrofauna benthic communities before, we aimed to check whether the same pattern occurs among meiofauna. A total of 12 higher meiofaunal taxa were recorded, with nematodes predominating at all stations. Non-parametric multivariate analyses demonstrated clear differences in meiofaunal abundance and composition between stations in the glacial bay and in the outer part of the fjord. Meiofaunal abundance increased with increasing distance from the source of disturbance, which in our study is tidal glaciers. Therefore, the current study demonstrates that the spatial structure of meiofauna is affected by the natural environmental disturbance, and analysis of meiofaunal assemblages can be used to assess the effect of such disturbances.     

Putting the meio- into stream ecology: current findings and future directions for lotic meiofaunal research

• 1 There is a paucity of research on epigean freshwater lotic meiofauna. This may result from a previous emphasis on interstitial (groundwater and hyporheic) meiofauna and/or a reliance on sampling methodologies in lotic systems which are inappropriate for meiofauna.
• 2 Meiofauna contribute much to the diversity of lotic ecosystems. Species lists for seven streams reveal that meiofauna contribute 58–82% of total species numbers, with rotifers and chironomids dominating most systems. The absence of taxonomic keys for most meiofaunal taxa in large areas of the world precludes a wider analysis of their contribution to lotic diversity and an assessment of biogeographical patterns and processes.
• 3 The trophic and functional role of meiofauna in lotic ecosystems is unclear. There are few estimates of meiofaunal production in freshwaters and biomass spectra have produced conflicting results for lotic meiofauna. Present static estimates suggest that the contribution of meiofauna to lotic productivity and biomass is small to moderate, but further studies incorporating a temporal component may provide a more realistic picture of the total contribution of meiofauna to biomass size spectra.
• 4 Meiofauna differ from macroinvertebrates in several respects apart from size and conceptual models for lotic ecosystems should include all metazoans if they are to be truly representative.
• 5 Information on the basic ecology of certain lotic meiofauna (i.e. nematodes, tardigrades, microturbellarians) is urgently required. For those groups whose distributional patterns are better understood (e.g. microcrustaceans), the mechanisms underpinning these patterns should be explored. It is essential that the importance of meiofauna is recognised by lotic ecologists; the only realistic way forward is for greater collaboration among meiofaunal ecologists and taxonomists and other lotic scientists.

     

Meiofauna distribution in Martel Inlet, King George Island (Antarctica): sediment features versus food availability

The meiofauna density in Martel Inlet, Admiralty Bay, King George Island (Antarctica) was studied in order to better understand the spatial patterns of meiofauna distribution in shallow polar seas. Sampling was undertaken by scuba-diving at 15 m depth at 7 stations during the summers of 1996/1997 and 1997/1998. The meiofauna was dominated by nematodes (>60%), followed by copepods, nauplii and polychaetes. The mean densities found during the 2 years studied (4.09 and 3.5᎒⁶ ind. m^-2) suggest that the shallow area of Martel Inlet is characterized by higher numbers of meiofaunal organisms than are found in other ecosystems, which is in agreement with other meiofaunal studies carried out in polar areas. It may be concluded that it is mainly the sedimentary features that influence the meiofauna distribution. However, food availability must contribute to the support of the higher meiofauna densities found in the inlet.     

The ecology of shallow water meiofauna in two New England estuaries

Summary The distribution and abundance of shallow subtidal meiofauna in four habitats in the Niantic and Pettaquamscutt estuaries in southeastern New England were studied from October 1964 through October 1965. Numbers of individuals ranged from 1,184 to 5,163/10 cm², and wet weights from 8.5 to 62.5 mg/10 cm². Nematodes were the dominant group, averaging 83% of the total numbers and 64% of the total biomass. Among the nematodes, epigrowth-feeding species were dominant at all four stations; in addition, deposit-feeding species were abundant at three stations where the amount of detritus was high relative to the fourth station. At these three stations ostracods and deposit-feeding polychaetes were also abundant.Nematodes showed marked seasonal changes i their species composition, with epigrowth-feeding species reaching maximum densities in spring and summer (coincident with observed increases in benthic microflora production), and deposit and omnivorous-feeding species in fall and winter (coincident with observed increases in organic detritus). Increases in the epigrowth feeders were responsible for significant increases in the total nematode populations in spring and summer.Ostracods were most abundant in late summer, fall and winter in association with the increases in detritus. Polychaetes, amphipods and lamellibranchs were most abundant in summer, the first-named in association with elevated water temperatures. Harpactacoid copepods were inconsistent in their seasonal distributions.Approximately 80% of the meiofauna occurred in the upper 3 cm of sediment. Copepods and ostracods were virtually limited to the upper 3 cm, while amphipods and lamellibranchs were limited to the upper 2 cm of sediment. Nematodes and polychaetes extended down to 5 cm (the lowest depth studied), although usually in significantly reduced numbers.Based on a thesis submitted in partial fulfillment of the requirements for the Ph.D. degree, University of Rhode Island, 1966. This study was supported in part by U.S. Bureau of Commercial Fisheries Grant No. 14-17-0007-189 (G). Some of the recent laboratory studies supported by National Science Foundation Grant No. GB 7796.     

Long-term nutrient enrichment elicits a weak density response by saltmarsh meiofauna

The effect of chronic nutrient enrichment on benthic meiofauna was examined in a whole-ecosystem experiment conducted in salt marshes in the Plum Island watershed of northeastern Massachusetts, USA. We compared abundances of total meiofauna, nematodes, copepods, ostracods, and annelids including Manayunkia aestuarina, in fertilized (where N and P was increased 15× in incoming tidal water throughout each growing season for 6 years) and reference marsh creeks to test for bottom-up responses. Although some responses to nutrient enrichment were evident, results did not match our expectations of strong increases in abundance. Variation in abundance between nutrient-enriched and reference creeks was detected in all three subhabitats but responses were inconsistent and variable over time, suggesting that natural variability was greater than variation induced by fertilization. Our results showed an overall weak negative correlation between meiofauna abundance and benthic microalgae (BMA) biomass partly because the BMA response to nutrient enrichment was relatively small and perhaps limited by grazing macrofauna and nekton. Our results suggest a better mechanistic understanding of the relationship between BMA and meiofaunal abundance is needed to fully understand how nutrient enrichment affects meiofauna.     

污水排海对小型底栖生物丰度和生物量的影响

为了解污水排海对小型底栖生物丰度和生物量的影响,于2011年4、8、10、12月对青岛汇泉湾第一海水浴场中潮带一个排污口附近不同距离站位的小型底栖生物进行了春、夏、秋、冬4个季度的采样调查.结果表明： 研究区域小型底栖生物年平均丰度为(1859.9±705.1) ind·10 cm^-2,最高值出现在距离排污口20和40 m的站位S₂和S₃,分别为(2444.9±1220.5)和(2492.2±1839.9) ind·10 cm^-2,最低值出现在距离排污口0 m的站位S₁,为(327.9±183.2) ind·10 cm^-2.小型底栖生物的年平均生物量为(1513.4±372.7) μg·10 cm^-2.小型底栖生物在丰度和生物量上呈现明显的季节变化,最高值出现在春季,最低值出现在夏季.共鉴定出11个小型底栖生物类群,包括线虫、桡足类、多毛类、寡毛类、缓步动物、海螨、涡虫、介形类、等足类、甲壳类幼体及其他类.自由生活海洋线虫是最优势的类群,占总丰度的83.1%,其次为底栖桡足类,占12.8%.在垂直分布上,小型底栖生物在0~2 cm表层分布最多,向深层呈现递减趋势,冬季部分向下迁移.Pearson相关分析表明,小型底栖生物丰度和生物量与沉积物中值粒径和有机质含量呈极显著负相关.此外,旅游等人为扰动也是影响小型底栖生物数量及分布的因素.与历史资料中的同类研究结果进行了比较,并探讨了线虫与桡足类丰度的比值
在有机质污染监测中的适用性.     

Distribution of meiofauna in Kongsfjorden,Spitsbergen

Kongsfjorden, a glacial fjord, is a typical fjord in the Spitsbergen (Svalbard archipelago) in the Arctic. The study supports a hypothesis that meiofauna and macrofauna are affected by natural environmental disturbances. Therefore, meiofaunal and macrofaunal analyses can be used to assess the effects of natural environmental disturbances in similar fjords in the Spitsbergen. Inputs from tidal glaciers create steep environmental gradients in sedimentation and salinity along the fjord. The magnitude of the glacial outflow diminishes towards the outer part of the fjord. Glacial-related physical stress causes reduced abundance, biomass and diversity among the meiofaunal assemblages in the inner part of the fjord. Based on quantitative and qualitative analyses of the composition of collected samples, three groups of meiofauna have been distinguished: one outer basin association and two in the inner, glacial bay. The presented results demonstrate that both the meiofauna and the macrofauna are affected on a similar scale by natural environmental disturbances. Therefore, as for macrofauna, meiofaunal analysis can be used to assess the effect of natural environmental disturbances.     

Intertidal meiofauna of a high-latitude glacial Arctic fjord (Kongsfjorden, Svalbard) with emphasis on the structure of free-living nematode communities

Meiofauna communities of four intertidal sites, two sheltered and two more exposed, in Kongsfjorden (Svalbard) were investigated in summer 2001 at two different tidal levels (i.e. the low-water line and close below the driftline, referred to as mid-water (MW) level). A total of seven meiofaunal higher taxa were recorded with nematodes, oligochaetes and turbellarians being numerically dominant. Mean total meiofaunal densities ranged between 50 ind. 10 cm⁻² and 903 ind. 10 cm⁻². Our data showed a clear decrease in total meiofaunal densities with increasing coarseness of the sediment. Total meiofaunal biomass varied from 0.2 g dwt m⁻² to 2 g dwt m⁻² and, in general, was high even at low meiofaunal densities, i.e. larger interstitial spaces in coarser sediments supported larger meiofauna, especially turbellarians. The results on the vertical distribution of meiofauna contrasted sharply with typical meiobenthic depth profiles on other beaches, probably in response to ice-scouring and concomitant salinity fluctuations. Oligochaetes were the most abundant taxon, with a peak density of 641 ind. 10 cm⁻² at Breoyane Island. They were mainly comprised of juvenile Enchytraeidae, which prohibited identification to species/genus level. Nematode densities ranged between 4 ind. 10 cm⁻² and 327 ind. 10 cm⁻². Nematodes were identified up to genus level and assigned to trophic guilds. In total, 28 nematode genera were identified. Oncholaimus and Theristus were the most abundant genera. The composition of the nematode community and a dominance of predators and deposit feeders were in agreement with results from other arctic and temperate beaches. Nematode genus diversity was higher at the more sheltered beaches than at the more exposed ones. Low-water level stations also tended to harbour a more diverse nematode communities than stations at the MW level. Differences in nematode community structure between low- and MW stations of single beaches were more pronounced than community differences between different beaches and were mainly related to resources quality and availability.     

The recovery of benthic foraminifera and bacteria after disturbance: experimental evidence

In experiments with living meiofauna, sediment is often sieved prior to incubation. This is primarily to remove macrofauna and to increase reproducibility among replicates. At the onset of the experiment, the bacteria are severely disturbed. The effects of these disturbances are ill-known but might affect the outcome of the experimental meiofaunal and biogeochemical studies substantially.We compared the disturbance induced by sieving with the disturbance in microcosms from which meiofauna was removed by flushing with argon. Both experimental situations were compared with untreated cores and the field situation. Neither sieving nor flushing induced changes in the composition of the foraminiferal community compared with the natural situation; the four most abundant species found in the field remained dominant during the experiment. Sieving led to a pronounced disturbance in both bacterial as well as foraminiferal abundance patterns. The depth distribution of some species seems to be related to food, although bacteria might play a regulating role as well.     

The response of meiofauna and microphytobenthos to engineering effects of fiddler crabs on a subtropical intertidal sandflat

Fiddler crabs are key bioturbators on tidal flats. During their intense bioturbation process, they manipulate large amounts of sediment, altering the physical state of existing materials. We investigated whether different types of sediment bioturbation produced by fiddler crabs modulate meiofaunal assemblages and microphytobenthic content. We hypothesized that sedimentary structures produced by burrowing (the burrow itself and the excavation pellets) and feeding (feeding pellets) generate different microenvironments compared with areas without apparent signs of fiddler crab disturbance, affecting both meiofauna and microphytobenthos, independent of the sampling period. Our results indicate that the engineering effects of burrow construction and maintenance and the engineering effects of fiddler crab foraging modulate meiofaunal assemblages in different ways. Overall, meiofauna from burrows and excavation pellets was more abundant and diverse than at control sites, whereas feeding pellets contained poor meiofaunal assemblages. By contrast, only foraging effects were detected on microphytobenthos; independent of the sampling period, Chl a and phaeopigment content were higher in the feeding pellets, but similar among burrows, excavation pellets and control sites. The present study demonstrates that the different engineering effects of fiddler crabs are an important source of habitat heterogeneity and a structuring agent of meiofaunal assemblages on subtropical tidal flats.     

Meiofauna of silty sediments in the coastal area of the North Adriatic,with special reference to sampling methods

At five coastal silty sediment stations ranging in depth from 8 to 30 m, the abundance and composition of meiofauna were investigated. Three methods of sampling were used, i.e. Pfleger corer, Van Veen grab and SCUBA divers. Four samples per station were taken. The mean density of total meiofauna was 660 ± 109 ind. 10 cm². The main meiofauna group was Nematoda, the second abundant was Copepoda, and third was Kinorhyncha. Statistical tests showed significant differences in meiofaunal abundance between corer and grab samples, and between corer and divers samples.Differences in meiofauna abundance between stations were found.     

On the Vertical Distribution of the Metazoan Meiofauna in Shelf Break and Upper Slope Habitats of the NE Atlantic

The metazoan meiofauna of nine stations in shelf break and upper slope areas (70 to 1500 m water depth) of the N.E. Atlantic were investigated in order to assess which environmental factors are important in the control of densities and sediment profiles. Total meiofaunal densities (ranging between 368 and 1523 ind/10 cm²) were correlated with bacterial densities, an important food source for meiofauna. However, considering sediment vertical distribution profiles, the relative importance of both food and oxygen on the meiofauna became obvious. A combination of both bacterial densities and oxygen supply could explain about 95% of the variability in the vertical profiles of the meiofauna densities. Meiofauna numbers increase in proportion to food availability in the surface sediment layers, but this relationship breaks down in deeper sediment layers where the oxygen supply is often limiting, particularly in fine sediments.