Transfer of lipids from plankton to blubber of harp and hooded seals off East Greenland

Sub-Arctic marine ecosystems are some of the most productive ecosystems in the world's oceans. The capacity of herbivorous zooplankton, such as Calanus, to biosynthesize and store large amounts of lipids during the short and intense spring bloom is a fundamental adaptation which facilitates the large production in these ecosystems. These energy-rich lipids are rapidly transferred through the food chain to Arctic seals. The fatty acids and stable isotopes from harp seal (Phoca groenlandica) and hooded seal (Cystophora cristata) off East Greenland as well as their potential prey, were analysed. The results were used to describe the lipid dynamics and energy transfer in parts of the East Greenland ecosystem. Even if the two seal species showed considerable overlap in diet and occurred at relatively similar trophic levels, the fatty acid profiles indicated that the bases of the food chains of harp and hooded seals were different. The fatty acids of harp seals originate from diatom-based food chain, whereas the fatty acids of hooded seals originate from dinoflagellate and the prymnesiophyte Phaeocystis pouchetii-based food chain. Stable isotope analyses showed that both species are true carnivores on the top of their food chains, with hooded seal being slightly higher on the food chain than harp seal.     

Trophic relationships along a bathymetric gradient at the deep-sea observatory HAUSGARTEN

Deep-seafloor communities, especially those from the ice-covered Arctic, are subject to severe food limitation as the amount of particulate organic matter (POM) from the surface is attenuated with increasing depth. Here, we use naturally occurring stable isotope tracers (δ¹⁵N) to broaden our rudimentary knowledge of food web structure and the response of benthic organisms to decreasing food supplies along the bathymetric transect (～1300–5600 m water depth) of the deep-sea observatory HAUSGARTEN. Encompassing five trophic levels, the HAUSGARTEN food web is among the longest indicating continuous recycling of organic material typical of food-limited deep-sea ecosystems. The δ¹⁵N signatures ranged from 3.0‰ for Foraminifera to 21.4‰ (±0.4) for starfish (Poraniomorpha tumida). The majority of organisms occupied the second and third trophic level. Demersal fish fed at the third trophic level, consistent with results from stomach contents analysis. There were significant differences in the δ¹⁵N signatures of different functional groups with highest δ¹⁵N values in predators/scavengers (13.2±0.2‰) followed by suspension feeders (11.2±0.2‰) and deposit feeders (10.2±0.3‰). Depth (=increasing food limitation) affected functional groups in different ways. While the isotopic signatures of predators/scavengers did not change, those of suspension feeders increased with depth, and the reverse was found for deposit feeders. In contrast to the results of other studies, the δ¹⁵N signatures in POM samples obtained below 800 m did not vary significantly with depth indicating that changes in δ¹⁵N values are unlikely to be responsible for the depth-related δ¹⁵N signature changes observed for benthic consumers. However, the δ¹⁵N signatures of sediments decreased with increasing depth, which also explains the decrease found for deposit feeders. Suspension feeders may rely increasingly on particles trickling down the HAUSGARTEN slope and carrying higher δ¹⁵N signatures than the decreasing POM supplies, which elevates the δ¹⁵N value of their tissues. Our results imply that a depth-stratified approach should be taken to avoid a misinterpretation of data obtained at different depths.     

Seasonal dynamics of sub-ice fauna below pack ice in the Arctic (Fram Strait)

Pronounced seasonality is a characteristic feature of polar ecosystems, but seasonal studies in the high-Arctic pack-ice zone are still scarce because of logistical constraints. During six expeditions (1994–2003) to the Fram Strait area between Greenland and Svalbard in winter, spring, early summer, late summer and autumn, the sub-ice habitat and fauna below the pack ice (0–1 m depth) were analyzed for seasonal patterns. Both environmental variables such as ice cover, temperature, salinity and chlorophyll a (chl a), as well as species composition, abundance and biomass of the sub-ice fauna showed distinct seasonal dynamics. Most species of the sub-ice fauna were found in early summer, followed by autumn, spring and late summer; the lowest number occurred in winter. The sub-ice fauna was dominated by copepod nauplii during all seasons. Next numerous was the small pelagic copepod Oithona similis, followed by occassional swarms of Pseudocalanus minutus and Calanus spp. Abundances of the sympagic fauna in the sub-ice water layer were much lower, with ectinosomatid copepods being usually the most numerous sympagic group. In the course of the year, total abundances of the sub-ice fauna showed a steep increase from the earliest sampling dates towards the end of winter/beginning of spring reaching maximum numbers then, and a decrease to minimum numbers in early summer. A second peak occurred in late summer, followed by a decrease towards autumn. This significant trend was due to the abundances of copepod nauplii and Oithona similis. Sympagic species were virtually absent during winter, and increased significantly in spring and early and late summer. A factor analysis revealed the variables ice cover and thickness, water temperature and salinity, as well as chl a as the major controlling factors for the seasonal patterns in different groups and species of the sub-ice fauna. Because of the special environmental conditions in the sub-ice habitat, and the unique species composition characterized by small taxa, young stages, and sympagic species, the seasonal dynamics of the Arctic sub-ice fauna differ substantially from those of the epipelagic zooplankton community in the Arctic Ocean.     

Trophic structure of vermetid reef community: High trophic diversity at small spatial scales

Stable isotopes were used to investigate contributions of autochthonous (i.e. benthic: epilithon and macroalgae) and allochthonous (i.e. pelagic: phytoplankton) organic matter sources to the diet of suspension-feeders, grazers and predators associated to small reef-pools (cuvettes) created by the reef-building species Dendropoma petraeum in the north-western coast of Sicily (Italy). Contributions of potential food sources were calculated using Bayesian mixing-models and integrated to a multivariate approach to highlight the diversity of C and N pathways within Dendropoma cuvettes. Both pelagic and benthic organic matter sources were exploited by benthic consumers, although clear differences were revealed in the various species depending on their feeding strategy. Three different trophic pathways were identified: one based mainly on phytoplankton, one based mainly on macroalgae and a third one mainly on epilithon. Suspension-feeders seemed to rely mainly on allochthonous organic matter sources, while grazers showed a wider diet spectrum. Predators revealed a high specialization in each of the three food chains and showed a distinct reliance on organic matter originated from benthic or pelagic sources. Stable isotopes evidenced here a marked differentiation of the trophic niche within the cuvette-associated community, which allows minimizing competition in very space-limited conditions.     

Trophic position of deep-sea fish—Assessment through fatty acid and stable isotope analyses

To investigate the trophic ecology of two of the dominant families of deep-sea fish (Macrouridae and Moridae) fatty acid and stable isotope analyses were applied to liver and muscle samples of five abundant species from the NE Atlantic. In conjunction with stomach content data these methods made it possible to identify differences in feeding strategies between the five study species as well as variation in feeding in relation to increasing depth and body size. Biomarkers identified strong similarities between Coryphaenoides armatus and Antimora rostrata though differences were found associating C. armatus more with the benthic food web whereas A. rostrata showed stronger links to the pelagic food web. While Lepidion eques was classified as a species linking benthic and benthopelagic food webs, both fatty acid and stable isotope data suggested that Coryphaenoides guentheri fed on an exclusively benthic diet. Coryphaenoides rupestris on the other hand were largely dependent on a copepod-based food web. Ontogenetic changes in feeding were found for both A. rostrata and C. armatus with the indication of a switch from active predation to scavenging occurring with increasing body size. Biomarkers also reflected the seasonal influx from the photic zone though changes were species-specific and probably reflected the variation in prey availability and abundance in response to these inputs. Our findings have thus demonstrated that the combined use of these biomarkers can elucidate trophic specialisations in situations where conventional methods alone previously provided insufficient data.     

A multiple-scale analysis of metazoan meiofaunal distribution in the deep Mediterranean Sea

In order to identify environmental factors driving the distribution and functioning of deep-sea fauna and the spatial scales of interactions, we carried out a multiple-scale investigation in the Mediterranean basin in which we compared two bathyal plains, located at the same depth (ca. 3000 m), but characterised by contrasting trophic conditions. We investigated meiofaunal abundance, biomass, community structure and biodiversity (expressed as richness of taxa) in relation to sediment characteristics, downward fluxes and food availability in the sediment. Samples were collected at all spatial scales (from small to macroscale) in two seasons. Our results indicated that deep-sea systems with different trophic conditions displayed different responses to the distribution of available energy and its spatio-temporal variability in the sediment. The analysis at a macroscale (>1000 km) indicated that meiofauna were controlled primarily by the trophic inputs to the deep-sea system. Spatial variability of meiofaunal parameters at a mesoscale (>50 km) was highest in the eastern Mediterranean and lowest in the western Mediterranean. Such differences are the consequence of the unpredictable inputs of organic matter in the oligotrophic eastern Mediterranean versus a more homogeneous distribution of food inputs in the mesotrophic western Mediterranean. At a smaller scale (local scale 7 km), in the western Mediterranean, the distribution of meiofaunal parameters was highly homogeneous, reflecting the homogeneous distribution of the food availability in the sediment. Our results indicated that the highly variable input and distribution of food sources in the deep eastern Mediterranean did not provide any “insurance” for the sustainability of the deep-sea faunal assemblages in the long term, thus leading to an uncoupling between resource availability and distribution of organisms. We conclude that the influence of energy availability on the deep-sea faunal distributions change at different spatial scales and that the analysis of spatial variability at mesoscales is crucial for understanding the relationships between deep-sea benthic fauna and environmental drivers.     

白令海和楚科奇海鱼类种类组成及其对生态环境变化的响应

根据第4次中国北极科学考察在白令海与楚科奇海进行的鱼类拖网调查资料,分析了白令海与楚科奇海鱼类生物的种类组成、优势种、物种多样性和区系特征,探讨了鱼类生物对北极气候快速变化的响应。结果表明,白令海与楚科奇海两个海域共鉴定鱼类生物14科41种;主要优势种类为粗壮拟庸鲽(Hippoglossoides robustus)、北鳕(Boreogadus saida)、短角床杜父鱼(Myoxocephalus scorpius)、斑鳍北鳚(Lumpenus fabricii)、粗糙钩杜父鱼(Artediellus scaber);从适温性来看,冷水性种类最多,有35种,冷温性种类6种;从栖息地生态类型来看,底层鱼类、近底层鱼类和中上层鱼类分别为35、5和1种;Shannon-Wiener多样性指数平均为1.21,呈现南高北低的特点,整体多样性水平不高;气候变化引起部分北极、亚北极海区鱼类出现不同程度的纬向和纵向移动,由此将引起北极渔业资源分布格局的变化。     

Composition of fish species in the Bering and Chukchi Seas and their responses to changes in the ecological environment

Based on trawl surveys in the Bering Sea and Chukchi Sea during the 2010 Chinese National Arctic Research Expedition, fish biodiversity characteristics, such as fish composition, dominant species, biodiversity, and faunal characteristics were conducted. We also discussed the responses of fishes to the quick changes in Arctic climate. The results showed that a total of 41 species in 14 families were recorded in these waters. The dominant species were Hippoglossoides robustus, Boreogadus saida, Myoxocephalus scorpius, Lumpenus fabricii, and Artediellus scaber. There were 35 coldwater species, accounting for 85.37%, and six cold temperate species, occupying 14.63%. The habitat types of fish could be grouped as follows: 35 species of demersal fishes, five benthopelagic fishes, and one pelagic fish. The Shannon–Wiener diversity index(H)(range between 0 and 2.18, 1.21 on average) was not high, and descended from south to north. Climate change has caused some fishes to shift along their latitudinal and longitudinal distribution around the Arctic and Subarctic areas, and this could lead to the decline of Arctic fishery resources.     

Ecosystem structure and resilience—A comparison between the Norwegian and the Barents Sea

Abundance and biomass of the most important fish species inhabited the Barents and Norwegian Sea ecosystems have shown considerable fluctuations over the last decades. These fluctuations connected with fishing pressure resulted in the trophic structure alterations of the ecosystems. Resilience and other theoretical concepts (top-down, wasp-waste and bottom-up control, trophic cascades) were viewed to examine different response of the Norwegian and Barents Sea ecosystems on disturbing forces. Differences in the trophic structure and functioning of Barents and Norwegian Sea ecosystems as well as factors that might influence the resilience of the marine ecosystems, including climatic fluctuation, variations in prey and predator species abundance, alterations in their regular migrations, and fishing exploitation were also considered. The trophic chain lengths in the deep Norwegian Sea are shorter, and energy transfer occurs mainly through the pelagic fish/invertebrates communities. The shallow Barents Sea is characterized by longer trophic chains, providing more energy flow into their benthic assemblages. The trophic mechanisms observed in the Norwegian Sea food webs dominated by the top-down control, i.e. the past removal of Norwegian Spring spawning followed by zooplankton development and intrusion of blue whiting and mackerel into the area. The wasp-waist response is shown to be the most pronounced effect in the Barents Sea, related to the position of capelin in the ecosystem; large fluctuations in the capelin abundance have been strengthened by intensive fishery. Closer links between ecological and fisheries sciences are needed to elaborate and test various food webs and multispecies models available.     

Food web structure of the coastal area adjacent to the Tagus estuary revealed by stable isotope analysis

The identification of energy sources, pathways and trophic linkages among organisms is crucial for the understanding of food web dynamics. Stable isotopes were used to identify the trophic level of food web components and track the incorporation of organic matter of different origins in the coastal ecosystem adjacent to the Tagus estuary. It was shown that the river Tagus is a major source of organic carbon to this system. Also, the wide difference in δ¹³C among the primary consumers allowed the identification of the pelagic and the benthic energy pathways. The maximum trophic level observed was 2.4 for Sepia officinalis. This value is indicative of a short food web. It was concluded that the diet of the upper trophic level species relies directly on the lower food web levels to a considerable extent, instead of relying mostly on intermediate trophic level species. Moreover, the δ¹⁵N values of primary consumers were very close to that of particulate organic matter, probably due to poorly known processes occurring at the basis of the food web. This lowers the trophic length of the whole food web. Reliance on benthic affinity prey was high for all upper trophic level secondary consumers.     

Feeding preferences of estuarine mysids Neomysis integer and Rhopalophthalmus tartessicus in a temperate estuary (Guadalquivir Estuary,SW Spain)

Mysid shrimps are an important component of estuarine food webs because they play a key role in energy transfer as intermediate prey. We investigated the seasonal, tidal and depth specific variation in the diet of the estuarine mysids Neomysis integer and Rhopalophthalmus tartessicus and explored its implications for the planktonic community structure of a temperate estuary (Guadalquivir Estuary, SW Spain). Neomysis integer is an opportunistic omnivore feeding mainly on mesozooplankton and on members of the detrital–microbial loop, shifting prey seasonally according to availability. In contrast, R. tartessicus showed a more carnivorous diet and shifted its target prey during seasons of low resource availability. Despite statistically significant differences in diet composition, both species shared prey of similar size, particularly juvenile Mesopodopsis slabberi, the most abundant mysid species in this estuary, and copepods. Although these similarities imply inter-specific resource competition, their co-existence is achieved by niche partitioning and spatial segregation: the higher osmoregulatory capacity and foraging plasticity of N. integer confers a broader niche breadth for this species allowing N. integer to inhabit the more stressful oligohaline region of the estuary where R. tartessicus cannot survive. We propose that this mechanism relaxes the potential for competition between N. integer and R. tartessicus.     

Trophic ecology of European sardine Sardina pilchardus and European anchovy Engraulis encrasicolus in the Bay of Biscay (north-east Atlantic) inferred from δ13C and δ15N values of fish and identified mesozooplanktonic organisms

European sardine (Sardina pilchardus) and European anchovy (Engraulis encrasicolus) are two species of economical and ecological significance in the Bay of Biscay (north-east Atlantic). However, the trophic ecology of both species is still poorly known in the area, and more generally, few studies have considered the potential trophic overlap between sardines and anchovies worldwide. This study aims to highlight the trophic links between the mesozooplankton and adults of these two pelagic fish in the Bay of Biscay, through carbon and nitrogen stable isotope analysis (SIA). Mesozooplankton and individuals of sardines and anchovies were collected during one season (spring 2010), over spatially contrasted stations within the study area. First, the potential effect of preservation (ethanol vs. freezing) and of delipidation (by cyclohexane) on mesozooplankton δ¹³C and δ¹⁵N values was assessed. Results demonstrated the necessity to correct for the preservation effect and for lipid contents in mesozooplankton for further analyses of sardines' and anchovies' diet through SIA. Next, this study highlighted the interest of working on identified mesozooplanktonic organisms instead of undetermined assemblages when unravelling food sources of planktivorous fish using stable isotopes. The inter-specific variability of isotope values within a planktonic assemblage was effectively high, probably depending on the various feeding behaviours that can occur among mesozooplankton species. Intra-specific variability was also significant and related to the spatial variations of baseline signatures in the area. To investigate the foraging areas and potential diet overlap of S. pilchardus and E. encrasicolus, mixing models (SIAR) were applied. Both fish species appeared to feed mainly in the neritic waters of the Bay of Biscay in spring and to select mainly small- to medium-sized copepods (e.g. Acartia sp., Temora sp.). However, E. encrasicolus showed a greater trophic plasticity by foraging more offshore and on a wider range of prey sizes, while S. pilchardus seemed more limited to coastal areas and the mesozooplanktonic species of these waters for feeding.     

Patterns of bathymetric distribution among deep-sea fauna at local spatial scale: comparison of mainland vs. insular areas

We have compared the distribution of mesopelagic, benthopelagic and benthic fauna between two areas: one on the continental side of the Catalan Sea (cCS: northwestern Mediterranean) and one to the SW of the Balearic Islands (SWB: southwestern Mediterranean) at depths between 147 and 2266 m. Based on 88 bathyal fish and crustaceans (Decapoda and Peracarida) dominant in these communities, we compared the maximum depth of occurrence (MDO) of (upper) middle-slope species and the minimum depth of occurrence (mDO) of lower-slope dwelling species. Mid-slope fish, decapods, peracarids and, within the latter, amphipods and cumaceans had a deeper MDO in the cCS than in the SWB. Depth differences between MDO of species were significant for all taxa, except isopods. In the same way, lower slope fish and decapods had a shallower mDO in the SWB than in the cCS. Within peracarids, the dominant taxon (amphipods) also followed this trend. Depth differences in mDO of species between the areas were significant for decapods and for amphipods (not for fish, nor all peracarids nor cumaceans). In summary, most taxa showed a deeper depth distribution of middle-slope species in the cCS, and a shallower depth distribution of lower-slope dwelling species in the SWB. This suggests that the whole community, from small detritus-feeders (peracarids) to top predators (fish) have a similar response to a common signal. Much basic information on the biology and possible environmental factors affecting deep-sea species distribution is not available, so causes of the trends demonstrated here cannot be fully evaluated. In spite of these obvious limitations, we have shown that (1) mesopelagic decapods (e.g., Gennadas elegans and Sergia robusta), with a higher dependence upon primary sources of food close to the surface primary production, showed greater differences in their mDO between the areas than benthopelagic (e.g., Acanthephyra eximia, Nematocarcinus exilis) and benthic (e.g., Stereomastis sculpta, Munida tenuimana, Geryon longipes) species, and (2) fish at lower trophic levels, deduced from fractional trophic levels, showed higher differences in the MDO than fish at higher trophic levels. Trophic position of species in food webs seems the most important factor affecting the distributional differences between contrasting areas.     

Difference of planktonic ciliate communities of the tropical West Pacific,the Bering Sea and the Arctic Ocean

Ciliates are important components in planktonic food webs,but our understanding of their community structures in different oceanic water masses is limited.We report pelagic ciliate community characteristics in three seas:the tropical West Pacific,the Bering Sea and the Arctic Ocean.Planktonic ciliate abundance had"bimodal-peak","surface-peak"and"DCM(deep chlorophyll a maximum layer)-peak"vertical distribution patterns in the tropical West Pacific,the Bering Sea and the Arctic Ocean,respectively.The abundance proportion of tintinnid to total ciliate in the Bering Sea(42.6%)was higher than both the tropical West Pacific(7.8%)and the Arctic Ocean(2.0%).The abundance proportion of small aloricate ciliates(10–20μm size-fraction)in the tropical West Pacific was highest in these three seas.The Arctic Ocean had higher abundance proportion of tintinnids in larger LOD(lorica oral diameter)size-class.Proportion of redundant species increased from the Arctic Ocean to the tropical West Pacific.Our result provided useful data to further understand ecology roles of planktonic ciliates in different marine habitats.     

Spatial organization of food webs along habitat gradients at deep-sea hydrothermal vents on Axial Volcano,Northeast Pacific

Deep-sea hydrothermal vents are characterized by steep spatial gradients and high temporal variability in habitat conditions. This leads to the organization of species distribution along spatial habitat gradients, which may constrain food resource utilization and food web structure. We conducted a stable-isotope-based study to test the hypothesis that food resource utilization is constrained by spatial habitat variability at diffuse hydrothermal vents on Axial Volcano, Northeast Pacific. Our study included the ten most biomass-prominent species and considered the temporal change in food web structure at recently created vent sites during three consecutive years. We related species average stable isotopic composition to their position between the center and the periphery of vent sites, using previously published data. Species spread widely along the δ¹³C axis, and showed a small variability in δ¹⁵N. This indicates that most species partition food resources between isotopically different carbon sources, and that they are not organized along predator–prey trophic chains. Particulate organic matter (POM) stable isotopic composition from a concomitant study corresponds to the signature of the expected diet for most organisms. Species average δ¹³C was significantly correlated to their relative position between the center and the periphery of vent sites. We relate this spatial variability in species isotopic composition to variability in the isotopic signature of both dissolved inorganic carbon (DIC) and POM. This spatial isotopic signal of consumers reveals the spatial structuring of food (POM) production and its consumption by the fauna. Accrual of species during the development of diffuse sites increased the inter-specific spread in δ¹³C, but did not increase the range in δ¹⁵N. Our results show that the spatial organization of species distribution results in a fragmented food web where species partition POM food resources according to their position in space. Shaping of species distribution by habitat gradients therefore constrains food web structure and the occurrence of predator–prey and competitive interactions.     

Interplay between top-down, bottom-up, and wasp-waist control in marine ecosystems

In October 2004, the North Pacific Marine Science Organization (PICES) sponsored a symposium to consider “Mechanisms that regulate North Pacific ecosystems: Bottom up, top down, or something else?” It sought to examine how marine populations, particularly the upper-trophic-level species, are regulated and to understand how energy flows through marine ecosystems. This introductory essay examines aspects of control mechanisms in pelagic marine ecosystems and some of the issues discussed during the symposium and in the 11 papers that were selected for this special issue. At global scales, the greatest biomass of fishes, seabirds and marine mammals tends to occur in regions of the world ocean with high primary production, e.g., the sub-arctic seas and up-welling regions of continental shelves. These large-scale animal distribution patterns are driven by food availability, not the absence of predators. At regional scales however, it is likely that current predation or past predation events have shaped local distributions, at least in marine birds and pinnipeds. Wasp-waist control occurs when one of the intermediate trophic levels is dominated by a single species, as occurs with small pelagic fishes of the world’s up-welling zones. Processes in these ecosystems may have features that result in a switch from bottom-up to top-down control.     

水层-底栖耦合生态动力学研究的某些进展

概述浅海生态系的水层系统与底栖系统耦合的基本原理,着重介绍有机质沉降动力学、底栖生态系统对有机质的响应、生物沉降和侧向平流、生物扰动和沉积物再悬浮研究的进展,结语中提出应予优先支持研究的科学问题。     

The Effect of Temporal Changes and Environmental Trophic Condition on the Isotopic Composition (ω13C and ω15N) of Atherina boyeri (Risso, 1810) and Gobius niger (L., 1758) in a Mediterranean Coastal Lagoon (Lake of Sabaudia): Implications for Food Web Structure

Abstract. ω¹³C and ω¹⁵N of organic matter sources and consumers were employed to analyse trophic differentiation between a benthic consumer, Gobius niger (L., 1758) (Pisces, Osteichthyes), and a pelagic consumer, Atherina boyeri (Risso, 1810) (Pisces, Osteichthyes) in a Mediterranean coastal lagoon (Lake of Sabaudia) in winter and summer 1999. Trophic differences between the two species throughout the two sampling periods were related to the environmental trophic condition (i. e. nutrient and phytopigment concentrations). Although these two fish have different habitats, they both exploited benthic organisms, above all in summer. When the nutrient and phytopigment concentrations were higher (summer), the most abundant benthic organisms were the main food sources for both species. In winter A. boyeri apparently shifted its feeding preferences to include zooplankton, confirming its opportunistic feeding strategy. Par-ticulate organic matter and algae seemed to play the main trophic role in the food web structure. Benthic invertebrates used both sources, while zooplankton seemed to rely exclusively on the particulate fraction. The overlapping and very depleted signatures of sedimentary and particulate organic matter highlights a strong link between sediments and water column, rarely observed in other aquatic ecosystems using stable isotopes. Such a finding further substantiates the importance of particulate organic matter as a food source in the study area.