Using biochemical and isotopic tracers to characterise organic matter sources and their incorporation into estuarine food webs (Rufiji delta,Tanzania)

Fatty acid biomarkers and stable isotope signatures were used to identify the sources of particulate and sedimentary organic matter and its input into the food web through the dominant consumer within the mangrove-dominated Rufiji estuary, Tanzania. Specific fatty acids were used to identify the preferred basal sources of dominant fauna (i.e. filter feeder bivalves, snails, crabs, shrimps, and three fish species), and their presence in the water and sediment samples in the estuary. Both fatty acid and stable isotope results revealed that food web in the Rufiji estuary depended on a variety of carbon sources (mangroves, allochthonous terrestrial inputs, macroalgae, and phytoplankton), contributing to a different degree into the diets of primary consumers and members of near-shore fish, but none of them were obligatory for the survival of these species. The δ¹⁵N values of major primary producers and consumers/predators revealed a trend for δ¹⁵N enrichment with increasing trophic level. The ratio of docosahexaenoic acid to eicosapentaenoic acid (DHA:EPA) decreased from pelagic to benthic feeding fish. This indicated that fish with different feeding modes derived their fatty acids from different primary sources of nutrition, and suggested that the DHA:EPA ratio may be a useful indicator of feeding mode.     

Role of grapsid crabs, Parasesarma erythrodactyla, in entry of mangrove leaves into an estuarine food web: a mesocosm study

The influence of the crab Parasesarma erythrodactyla on the entry of the organic matter derived from Avicennia marina mangrove leaves in a sub-tropical mangrove ecosystem of southeast Queensland, Australia, was simulated using tidal mesocosms. Degradation of mangrove leaf organic matter was followed by analysing the fatty acid composition, carbon, and nitrogen isotopic signatures of the surface sediment and suspended particulate organic matter (SPOM) with and without the presence of crabs. Assimilation of mangrove organic matter by P. erythrodactyla was also assessed by stable isotope and fatty acid analyses in tissues and faeces. Results of the chemical tracer analyses question the adaptability of P. erythrodactyla to a diet comprised exclusively of mangrove leaves, and suggest that these organisms were dependent on additional food sources in their natural environment. Crab processing of senescent leaves significantly accelerated the transfer of mangrove organic matter to the surface sediments, as shown by a higher C/N ratio, a higher contribution of long-chain fatty acids and a more depleted C isotopic signature of sediment samples in the mesocosms with crabs compared to those without crabs.     

Sea ice microbial production supports Ross Sea benthic communities: influence of a small but stable subsidy

Diversity in guilds of primary producers enhances temporal stability in provision of organic matter to consumers. In the Antarctic ecosystem, where temporal variability in phytoplankton production is high, sea ice contains a diatom and microbial community (SIMCO) that represents a pool of organic matter that is seasonally more consistent, although of relatively small magnitude. The fate of organic material produced by SIMCO in Antarctica is largely unknown but may represent an important link between sea ice dynamics and secondary production in nearshore food webs. We used whole tissue and compound-specific stable isotope analysis of consumers to test whether the sea ice microbial community is an important source of organic matter supporting nearshore communities in the Ross Sea. We found distinct gradients in delta13C and delta15N of SIMCO corresponding to differences in inorganic carbon and nitrogen acquisition among sites with different sea ice extent and persistence. Mass balance analysis of a suite of consumers demonstrated large fluxes of SIMCO into the nearshore food web, ranging from 5% to 100% of organic matter supplied to benthic species, and 0-10% of organic matter to upper water column or pelagic inhabitants. A delta13C analysis of nine fatty acids including two key biomarkers for diatoms, eicosapentaenoic acid (EPA, 20:5omega3), and docosahexaenoic acid (DHA, 22:6omega3), confirmed these patterns. We observed clear patterns in delta13C of fatty acids that are enriched in 13C for species that acquire a large fraction of their nutrition from SIMCO. These data demonstrate the key role of SIMCO in ecosystem functioning in Antarctica and strong linkages between sea ice extent and nearshore secondary productivity. While SIMCO provides a stabilizing subsidy of organic matter, changes to sea ice coverage associated with climate change would directly affect secondary production and stability of benthic food webs in Antarctica.     

Feeding strategies of some demersal fishes of the continental slope and rise off the Mid-Atlantic Coast of the USA

Stomach contents of 729 fishes comprising 16 species were examined from the continental slope and rise off the Middle Atlantic States of the USA. Two main feeding modes among demersal deep-sea fishes were evident: those feeding primarily on pelagic food items, and those feeding on benthic invertebrates. Both pelagic and benthic predators were euryphagous. Most pelagic predators also fed on the epibenthos. These findings support Dayton and Hessler's (1972) suggestion that benthic predators should have a generalized diet which may be responsible for the high diversity found in the deep-sea infauna. The mesopelagic fauna is an important food source for some demersal fishes on the continental slope. Pelagic prey, which are also important to ecologically dominant demersal fishes on the lower slope and continental rise, may be nutritionally supported by suspended particulate organic matter in a nepheloid layer close to the bottom, and they may exist in much higher concentrations than in the bathypelagic zone above.Virginia Institute of Marine Science Contribution No. 835.     

Biochemical composition of sedimentary organic matter in the coral reefs of Lakshadweep Archipelago,Indian Ocean

Surface sediments were collected from the shore and lagoons of Kavaratti, Kadamat Agatti and Pitti islands of Lakshadweep Archipelago during May 2015 and analysed for biochemical composition and quality of organic matter. The biochemical composition of sedimentary organic matter from the entire study area was characterised by the dominance of carbohydrates (CHO) followed by proteins (PRT) and finally lipids (LPD). PRT:CHO ratios were less than 1 and indicated the presence of aged organic matter in the islands. The poor nutritional quality of sediments to support benthic fauna was evident from the values of LPD:CHO ratios. The refractory nature of sediments and less availability of food to benthic source was supported by BPC:TOC ratios. Based on estimated ratios and biopolymeric carbon values, the trophic status of the study area was categorised as oligotrophic.     

Hagfish in the New Zealand fjords are supported by chemoautotrophy of forest carbon

Forest litter is often considered to be a minor energy source to marine communities due to its refractory nature. Large volumes of forest litter are deposited in the New Zealand fjords, and likely recycled into available energy by microbial activity. In this study we used evidence from stable isotope analyses to test whether recycled carbon from chemoautotrophs was an important contributor to the diet of hagfish (Eptatretus cirrhatus). We then analyzed fatty acid biomarkers from the chemoautotrophic clam Solemya parkinsoni and E. cirrhatus to further discriminate the contribution of marine, terrestrial, and chemoautotrophic sources. Bulk isotopic signatures of E. cirrhatus varied considerably (delta13C, from -29.2 per thousand to -16.7 per thousand; delta15N, from -2.8 per thousand to +15.5 per thousand; delta34S, from -21.7 per thousand to +16.7 per thousand) and indicated that a significant percentage of organic matter (38-51%) originated from chemoautotrophs (delta13C, -31.3 per thousand +/- 0.1 per thousand [mean +/- SE]; delta15N, -5.7 per thousand +/- 0.2 per thousand; delta34S, -32.per thousand +/- 3.8 per thousand). Fatty acid biomarkers were depleted in 13C, particularly cis-vaccenic acid (18:1omega7: delta13C, -39.0 per thousand) indicating specific microbial origins of carbon. A high proportion of forest litter in sediments, coupled with isotopic and fatty acid biomarker results, indicates that terrestrial organic matter is a dominant contributor to this marine benthic system. This study demonstrates a clear linkage between terrestrial and marine ecological processes.     

Sources and major biogeochemical pathways of organic matter in the mangrove system of Rufiji estuary

Fatty acids were extracted from the surface sediments (10 cm) of three sampling sites of Rufiji estuary to infer their sources and biogeochemical pathways. The fatty acids ranging from C₈ to C₂₄ were distinguished from this study, and were broadly classified into saturated (SFAs), monounsaturated (MUFAs) and polyunsaturated fatty acids (PUFAs). SFAs were found to be the major fraction at station 1 and 3 where as at station 2, MUFAs dominated. A total of 19 fatty acids with a total concentration of 64.91 μg/g dry weight were characterised at station 1. C_16:0 was the most abundant fatty acid contributing 21.94% of total fatty acids (TFAs). C_22:2 was the second most abundant, which accounted for 9.46% of TFAs. Fatty acids ranging from C₁₂ to C₂₄ were identified at station 2. C₂₀ fatty acid was the most abundant fatty acid contributing 21.94%, followed by C_16:0. At station 3, fatty acids ranging from C₈ to C₂₄ were obtained. The PUFA C_20:5n-3 was the most abundant fatty acid contributing 21.65%, followed by C_24:0 (15.00% of TFAs). The ratio of lower molecular weight (LMW) to higher molecular weight (HMW) biomarkers was used as an indicators to distinguish higher plants organic matter from algae-derived fatty acids.     

Coral cavity sponges depend on reef-derived food resources: stable isotope and fatty acid constraints

The diet of cavity sponges on the narrow fringing reefs of Curaçao, Caribbean was studied. The origin and resources of the bulk food of these sponges, i.e., dissolved organic matter (DOM), were identified using stable carbon and nitrogen isotopes and fatty acid biomarkers. We found that phytoplankton and its derived DOM from the adjacent open sea and from reef overlying water is not the main source of food for most of the sponges examined nor is bacterioplankton. Interestingly, dual stable isotope signatures (δ¹³C_org, δ¹⁵N_org) and fatty acid biomarkers appoint coral mucus and organic matter derived from crustose coralline algae (CCA) as probable food sources for encrusting sponges. Mucus-derived DOM may contribute up to 66% to the diet of examined sponges based on results of dual isotope mixing model analysis. The contribution of CCA (as purported representative for benthic algae) was smaller with values up to 31%. Together, mucus- and CCA-derived substrates contributed for 48–73% to the diet of sponges. The presence of the exogenous fatty acid 20:4ω6 in sponges, which is abundant in coral mucus of Madracis mirabilis and in CCA, highlights these reef-derived resources as sources of nutrition for DOM feeding cavity sponges. The relatively high concentrations of exogenous 20:4ω6 in all sponges examined supports our hypothesis that the bulk of the food of the cavity sponge community is reef-derived. Our results imply that cavity sponges play an important role in conserving food and energy produced within the reef.     

Organic matter composition in sediments of the Oman Margin

The quantity and biochemical composition of organic matter (OM) in the sediments underlying and below oxygen minimum zone (OMZ) in the Arabian Sea were studied to provide information on the diagenetic processes of organic carbon under different environmental conditions. Concentrations of total organic carbon (TOC), total nitrogen (TN) and total hydrolysable amino acids (THAA) were significantly higher in sediments within rather than below OMZ, while those of total carbohydrates (TCHO) were slightly lower in the latter, suggesting the presence of a larger supply of labile compounds into the sea bottom at the shallower site (i.e. within OMZ). Hydrolysable amino acid and carbohydrate contribution to TOC were even lower (about 10% in surficial sediments) than the values obtained from the abyssal oligotrophic North Pacific, suggesting that OM food availability in the Oman Margin sediments within OMZ was lower than that observed at abyssal depths. The presence of the highest THAA and TCHO concentrations in the top 40 mm of the sediment core at both sites reflected the presence of bioturbation processes. In contrast with the general view of the deep sea as a stable and constant system, below OMZ in the Arabian Sea sediments, some differences were observed in the two investigated cores indicating the presence of a certain spatial variability in OM content and diagenesis.     

Organic matter composition in sediments of the Oman Margin

The quantity and biochemical composition of organic matter (OM) in the sediments underlying and below oxygen minimum zone (OMZ) in the Arabian Sea were studied to provide information on the diagenetic processes of organic carbon under different environmental conditions. Concentrations of total organic carbon (TOC), total nitrogen (TN) and total hydrolysable amino acids (THAA) were significantly higher in sediments within rather than below OMZ, while those of total carbohydrates (TCHO) were slightly lower in the latter, suggesting the presence of a larger supply of labile compounds into the sea bottom at the shallower site (i.e. within OMZ). Hydrolysable amino acid and carbohydrate contribution to TOC were even lower (about 10% in surficial sediments) than the values obtained from the abyssal oligotrophic North Pacific, suggesting that OM food availability in the Oman Margin sediments within OMZ was lower than that observed at abyssal depths. The presence of the highest THAA and TCHO concentrations in the top 40?mm of the sediment core at both sites reflected the presence of bioturbation processes. In contrast with the general view of the deep sea as a stable and constant system, below OMZ in the Arabian Sea sediments, some differences were observed in the two investigated cores indicating the presence of a certain spatial variability in OM content and diagenesis.     

Meiobenthos distribution pattern in the tropical East Atlantic: indication for fractionated sedimentation of organic matter to the sea floor?

To test the extent to which sea surface productivity governs the distribution pattern of benthic organisms, meiobenthic standing stocks were investigated on the shelf, continental margin and the adjacent abyssal plains off the western African coast between Guinea (10°N) and Angola (18°S). The area of investigation is characterized by gradients in surface productivity due to spatially and seasonally varying coastal upwelling. Reflecting the dependency of deep-sea organisms on organic matter input from the euphotic zone, similar gradients ought to be expected within the benthos. Meiofaunal abundances and biomasses (including Foraminifera) from a total of 57 stations along 13 transects across the continental margin showed a fairly close correlation with sediment-bound chloroplastic pigment concentrations, indicating the sedimentation of particulate organic matter from phytoplankton production. However, certain discrepancies in faunal and pigment distribution patterns were found in regions apart from the centres of enhanced primary␣productivity, i.e. apart from the upwelling centres: whereas pigment concentrations in the sediments were still comparably high, meiofaunal numbers in those peripheral areas were generally lower. It is suggested that smaller/lighter phytodetritial matter, transported over long distances by subsurface currents and exposed to ongoing microbial degradation during its passage, probably does not have the same energy content as the relatively fast-sinking, larger phytodetritus aggregates (“marine snow”), which are found in centres of enhanced primary productivity and support extensive benthic stocks. Thus, meiobenthic abundances in relation to sediment-bound pigment concentrations on the western African continental margin may indicate fractionated sedimentation of organic matter to the sea floor. Received: 28 May 1997 / Accepted: 14 July 1997     

Distribution and behaviour of polycyclic aromatic hydrocarbons in subtropical river estuary,Okinawa, Japan

Surface water samples were collected along the salinity gradient of Manko estuary in Okinawa, Japan, between January and November 2012, to examine spatial and temporal variability in the distribution, composition, behaviour, and sources of particulate polycyclic aromatic hydrocarbons (PAHs). Concentrations of total particulate PAHs ranged from 169.7?ng?g^?1 at riverine station (N1) in November 2012 to 922?ng?g^?1 at another riverine station (K3) in May 2012. Based on observed environmental behaviour, particularly relationships to organic matter, the PAHs appear divisible into two groups: group I (low molecular weight) PAHs were characterised by low particles affinity and strong correlations with a pool of autochthonous organic matter (e.g. high concentrations of polyunsaturated fatty acids), reflecting processes controlling their delivery to the estuary; and, in contrast, group II (high molecular weight) PAHs were characterised by high particles affinity and exhibited estuarine distribution primarily controlled by suspended solid load and inputs from terrigenous sources through river inflows. This relationship, coupled with isomer ratios indicative of combined mixed sources from petroleum and petroleum combustions, indicates that terrestrial run-off is the dominant mode of delivery of particulate PAHs in the estuary. The relationships between distinct groups of PAHs and lipid biomarkers demonstrated in this study suggest that very specific pools of particulate organic matter, as opposed to concentrations of particulate organic carbon alone, can be important in the transport and fate of hydrophobic contaminants.     

Suspended particulate organic matter in a Mediterranean submarine cave

Submarine caves display a paucity of benthic density and biomass that may be related to low trophic resources. Analysis of organic carbon, organic nitrogen, carbohydrate, protein and lipid content of suspended particulate matter was made during the period July 1985–July 1987 in a Mediterranean cave (Marseille, France) in order to determine any differences in the particulate organic matter composition along an horizontal transect. Particulate organic matter content clearly declined from the entrance of the cave to the dark inner area. This impoverishment could not be explained by a simple decrease in a few organic compounds, but appeared to be related to the combination of a decrease in both the amount and the composition of the suspended particles. Three progressive levels of impoverishment were identified towards the dark inner area of the cave: (i) decreasing amounts of seston; (ii) decreasing organic content of particles; (iii) increasing proportions of the geopolymeric (i.e., humic) components in the remaining organic matter, indicating increased degradation. The cave appeared to be sharply divided into two distinct sections — a twilight outer section whose waters contained slightly lower amounts of particulate organic matter than the open sea, and a dark inner section, 8 to 10 m higher, separated from the outer section by a steep rise and with waters of low organic matter content. The water in the twilight section appeared to be in thermal equilibrium with the open sea, and that in the dark inner section displayed thermal stratification. These differences indicated the presence of two distinct water bodies with contrasting average residence times, estimated as 1 d in the outer twilight section and 8 d in the dark inner section. The joint action of sedimentation and degradation resulted in an abrupt depletion of particulate organic matter in the dark inner section accompanied by a decrease in the benthic fauna. The decline in benthic heterotroph populations is probably related to the abrupt transition to oligotrophic conditions.     

Trace Organic Matter in the Ria Formosa,Portugal

The Ria Formosa lagoon, Portugal, receives organic matter from many natural and anthropogenic sources. This paper presents an overview of the contribution these sources made through the use of lipid biomarkers. Sixty-one surface sediment samples were collected and analysed for a range of lipids. Sewage materials were confined to regions within 2 km of discharge points; phytoplankton biomarkers (sterols and fatty acids) suggest production was greatest in the inner parts of the lagoon that also have greatest inorganic nutrient enrichment; terrestrial organic matter was present in relatively low concentrations as shown by both sterol and fatty alcohols; bacteria were widespread although the Sulphate Reducing Bacteria (SRBs, shown by the presence of 3-OH fatty acids) were located near but not adjacent to sewage discharge points.     

Trace Organic Matter in the Ria Formosa, Portugal

The Ria Formosa lagoon, Portugal, receives organic matter from many natural and anthropogenic sources. This paper presents an overview of the contribution these sources made through the use of lipid biomarkers. Sixty-one surface sediment samples were collected and analysed for a range of lipids. Sewage materials were confined to regions within 2 km of discharge points; phytoplankton biomarkers (sterols and fatty acids) suggest production was greatest in the inner parts of the lagoon that also have greatest inorganic nutrient enrichment; terrestrial organic matter was present in relatively low concentrations as shown by both sterol and fatty alcohols; bacteria were widespread although the Sulphate Reducing Bacteria (SRBs, shown by the presence of 3-OH fatty acids) were located near but not adjacent to sewage discharge points.     

Is benthic trophic structure affected by food quality? The Dogger Bank example

The aim of this study was to evaluate the relationship between food quality and trophic structure of macrobenthic communities of the Dogger Bank. Therefore, 21 stations along different depth gradients (a transect across the central Dogger Bank and at a station cross in the north-eastern part) were sampled and analysed for macrofauna trophic structure and food supply, indicated by phytopigment composition and organic matter contents of the sediment. The study revealed differences in the trophic structure of the benthic community in areas with contrasting food supply. The trophic structure reflected differences in relative quality of organic matter. The sandy sediments on top of the bank receiving intermediate quantities of high food quality were inhabited by high abundances of interface feeders and sand-licking amphipods of the genus Bathyporeia. The southern stations characterised by a comparatively high amount of intermediate food quality were dominated by interface feeders such as Amphiura filiformis. The northern and north-eastern stations showing the highest diversity of feeding types were characterised by a comparatively low, but highly variable input of intermediate-quality organic matter consisting of both fresh and degraded organic matter. Results are discussed with a special emphasis on specific Dogger Bank features such as benthic primary production and frontal system characteristics.Communicated by O. Kinne, Oldendorf/Luhe     

Biodiversity and trophic structure of nematode assemblages in seagrass systems: evidence for a coupling with changes in food availability

Nematode assemblage composition, trophic structure and biodiversity were followed over an annual cycle in a Posidonia oceanica bed of the NW Mediterranean to test the response to temporally changing food availability (sediment organic matter, bacterial and microphytobenthic biomass). The sediment-water interface of the seagrass meadow was characterised by high particulate organic matter concentrations. Also, seagrass sediments had high organic matter content, chloropigment concentrations and bacterial biomass. All organic matter components (i.e. carbohydrates, proteins and lipids) changed temporally, with higher concentrations in winter and spring and lower concentrations in summer; however, overall, large amounts of organic compounds were potentially available for (seagrass) benthic consumers throughout the year. Nematode assemblage in the P. oceanica bed maintained a high genus number (88 genera), and a trophic structure tightly coupled with the composition of the potential food sources. In agreement with the relevance of microphytobenthos, epigrowth-feeder nematodes (2A) represented the dominant trophic guild. The biomass of predator nematodes (2B) was significantly correlated with the biomass of other nematodes. Non-selective deposit feeders (1B) displayed a close relationship with the concentration of proteins, carbohydrates and biopolymeric carbon (i.e. labile organic detritus). Conversely to what was expected, epigrowth feeders were not correlated with chloropigment concentrations, but showed a significant relationship with the number of dividing bacteria, possibly suggesting a trophic plasticity of the 2A feeding guild. The coupling between temporal changes in food sources and temporal variability of the nematode trophic structure was highlighted by a CANOCO analysis, which allowed us to identify and associate, at each sampling time, nematode genera and their feeding habits, with specific environmental variables and food indicators. Furthermore, species diversity (H') and evenness (J) calculated on nematodes identified to the genus level displayed evident temporal changes, also reflected by the index of trophic diversity (ITD). Both structural and functional diversity were coupled with high concentration and highly heterogeneous composition of the food sources (including organic detritus, microphytobenthic algae and bacteria) potentially available to nematodes. These data suggest that temporal changes in quantity and quality of food sources do not only influence nematode dynamics and trophic composition, but also influence nematode structural and functional diversity.     

Variability of particulate organic carbon and nitrogen in the Namibian upwelling system

Particulate organic carbon (POC) and nitrogen (PON) were analyzed during cruises undertaken in September 1985 and April 1986 along the Namibian upwelling system. The main objectives were to provide estimates of standing stocks of particulate organic matter (POM) and analyze its temporal and spatial variability. Average estimates of total carbon standing stock (0 to 100 m depth) accounted for 1.2 g-at C m^–2 during active and 1.32 g-at C m^–2 during abated upwelling. Zooplankton carbon was estimated as 0.22 and 0.27 g-at C m^–2, respectively, for both periods. POM was generally concentrated near the surface, especially during abated upwelling. During abated upwelling, POM was not only abundant inshore but also over the shelf, as a response of a more diffuse frontal system and a more strongly stratified water column. Cross-shelf gradients were more significant during active upwelling, while alongshore gradients accounted for most of the variance of particulate organic matter during abated periods. This result was a consequence of the seasonal intrusion of warm, Angolan water from the north during the period of minimum upwelling, and resulted in poorer POM concentrations and higher consumer: producer ratios (24.4%). Nevertheless, this last conclusion should be regarded with caution due to the lack of comparative interannual variability. A 48 h study at a fixed station permitted analysis of the daily variability in POM during the intrusion process. Changes in the thickness of the surface mixed layer due to irregular time-spaced pulses of non-homogeneous water masses resulted in sudden enrichments and renewals of phytoplankton and zooplankton populations in a matter of hours.     

Eutrophication and trophic structure in response to the presence of the eelgrass <Emphasis Type="Italic">Zostera noltii</Emphasis>

In estuaries, eelgrass meadows contribute to fundamental ecosystem functions of estuaries, providing food to several predators and buffering the negative effects of eutrophication. We asked whether the presence of the eelgrass Zostera noltii decreased the nitrogen concentration in the overlying water, affected the sources of nitrogen sequestrated by primary producers and changed the benthic and pelagic food web structures. We also studied the importance of these food webs in providing food to fish. We compared bare sediment to sediment covered by a Z. noltii meadow, and examined nutrient concentrations in the water column and δ¹⁵N in primary producers as indicators of anthropogenic inputs of nutrients. We then measured both δ¹³C and δ¹⁵N in the tissues of plants and consumers to establish food web structures. There were no differences in the concentrations and sources of nitrogen between sites. Rather, δ¹⁵N values indicated anthropogenic inputs of N (e.g. sewage discharges, agriculture) in both sites. There were no major differences in the structure of the planktonic food web, which was in part sustained by particulate organic matter and supported most predator fish, and in the structure of the benthic food web. Nonetheless, there were differences in the sources of food for omnivore consumers and for the detritivore Scrobicularia plana. Overall, the benthic food web did not use food derived from the eelgrass or macroalgae deposited on the substratum. Suspension feeders used particulate and sediment organic matter, whereas the δ¹³C and δ¹⁵N values of the other consumers indicated a likely contribution of benthic microalgae. Furthermore, in both habitats we found large variability in the isotope signatures of benthic macrofauna consumers, which did not allow distinguishing clearly different trophic groups and indicated a high level of omnivory and a mixed diet opportunistically making use of the availability of food in the surroundings.