Reproduction of Calanus glacialis in the Laptev Sea, Arctic Ocean

Abundance and reproductive biology (gonad maturation and egg production) of the Arctic copepod Calanus glacialis were studied in the Laptev Sea and adjacent Arctic Ocean in September 1993 and from July to September 1995. Both abundance and reproductive activity were subject to strong spatial and seasonal variability, which was related to the ice cover, feeding conditions and circulation pattern. Maximum abundance of the C. glacialis population was generally confined to the outer shelf and slope with depths between 50 and 1000 m. During both cruises, highest egg production rates and largest number of young copepodite stages were observed in the eastern Laptev Sea, where the development of the C. glacialis population seems to follow the opening of the “Siberian Polynya”. In the western part, which is usually covered by pack ice, females were all immature, and no young stages were found. However, females responded quickly to a temporary opening of the ice there in 1995 and spawned. Starvation experiments showed that food-independent reproduction fuelled by internal energy resources was at least partly responsible for relatively high egg production rates at low ambient food concentrations. Egg production rates in starved females were considerably higher than those previously reported. Accepted: 14 June 2000     

Mesozooplankton assemblages in the shallow Arctic Laptev Sea in summer 1993 and autumn 1995

Mesozooplankton distribution and composition in the very shallow part of the Siberian Laptev Sea shelf were studied during the German-Russian expeditions “Transdrift I” (August/September 1993) and “Transdrift III” (October 1995). Maximum abundances were found close to the outflow of the Lena River (7,965 ind. m⁻³) and in the Yana river mouth (38,163 ind. m⁻³). Lowest abundances occurred in the northeast and west of the Laptev Sea (64–95 ind. m⁻³). Highest biomass values (104–146 mg DM m⁻³) were determined in the northern and northeastern part of the shallow Laptev Sea, as well as close to the river outflows, with a record biomass maximum in the Yana river mouth (270 mg DM m⁻³). Biomass minima were situated north of the Lena Delta and in the western part of the shallow Laptev Sea (0.3–1.0 mg DM m⁻³). Copepods dominated in terms of abundance and biomass. Cluster analyses separated four mesozooplankton assemblages: the assemblage “Lena/Yana” in the southern part, “Eastern-central” in the centre, “Kotelnyy” in the eastern part and “Taimyr” in the western part of the shallow Laptev Sea. The small-sized neritic and brackish-water copepods Drepanopus bungei, Limnocalanus grimaldii and Pseudocalanus major occurred in enormous numbers and made up the bulk of zooplankton abundance and biomass in the very shallow part of the Laptev Sea close to the rivers Lena and Yana. In the more northern and northeastern areas, Calanus glacialis, P. minutus and P. major were dominant copepod species, whereas Oithona similis and Acartia sp. became important in the western Laptev Sea. Appendicularians, as well as hydromedusae and the chaetognath Sagitta sp., contributed significantly to abundance and biomass, respectively, but not over the entire area studied. One can identify taxon-specific distribution patterns (e.g. Sagitta predominated the biomass in a zone between the area heavily influenced by Lena/Yana and the offshore area to the north), which differ from the patterns revealed by cluster analysis. Hydrographic features, especially the enormous freshwater inflow, apparently determine the occurrence and formation of zooplankton aggregations. Extremely high numbers of small-sized neritic and brackish-water copepods occurred locally, which were probably also supported by excellent feeding conditions.     

Transfer function between surface sediment diatom assemblages and sea-surface temperature and salinity of the Labrador Sea

Q-mode factor analysis (CABFAC) of 38 diatom species in 66 surface sediment samples from the Labrador Sea allowed the definition of six assemblages explaining 95.69% of the total variance. The first assemblage (40.12% of the total variance) is represented by Thalassiothrix longissima, a species characteristic of the Irminger Current and associated with the West Greenland Current. The second assemblage (34.52% of the total variance) is represented by Thalassiosira gravida (resting spores), a cold water species associated with the Labrador Current. A third assemblage (11.43% of the variance) represented by Chaetoceros spp. is concentrated in the northeastern section of the Labrador Sea and associated with upwelling and high productivity. The fourth, the fifth and the sixth assemblages, represented, respectively, by Coscinodiscus marginatus, Coscinodiscus oculus iridis, and Coscinodiscus divisus, and Nitzschia frigida, are less significant and account, respectively, for 4.95, 2.62 and 2.04% of the total variance. Transfer functions derived by relating factor assemblages to surface water temperature and salinity (August and February) give standard errors of estimate of ±0.93°C (August) and ±0.64°C (February) for temperature, and of ±0.58‰ (August) and ±0.45‰ (February) for salinity. High correlations between the measured and estimated parameters confirm the validity of the paleoecological equations for the reconstruction of the Labrador Sea paleoclimate.     

The sub-ice fauna of the Laptev Sea and the adjacent Arctic Ocean in summer 1995

The sub-ice habitat and fauna in the Laptev Sea and the adjacent Arctic Ocean were investigated during the “Polarstern” cruise ARK XI/1 in summer 1995. At the ice-water interface a thin thermo- and halocline developed at many stations due to melting processes. In the lower centi- to decimetres of the ice, an accumulation of organic matter was found (particulate organic carbon: 1.9 mg l⁻¹, chl a: 3.3 μg l⁻¹) that may have provided a food source for the fauna. The water layer directly beneath the ice was inhabited by high numbers of various nauplii (130–23911 ind. m⁻³), and two ecological groups, the pelagic sub-ice fauna that originates from the surface water plankton, and the sympagic sub-ice fauna that migrates into this boundary layer from the ice interior. The pelagic fauna dominated the sub-ice community both in terms of species number and abundance. Both groups mainly comprised small copepods (e.g. Oithona similis, Oncaea borealis, Pseudocalanus spp., Halectinosoma spp., Tisbe spp.), but foraminifers and pteropods, for example, also occurred regularly. Diversity was generally low. Factors influencing the composition and abundance of the sub-ice fauna were most likely water depth, salinity and sea-ice sediments. Accepted: 6 July 1998     

Distribution of diatom surface sediment assemblages within Effingham Inlet, a temperate fjord on the west coast of Vancouver Island (Canada)

Twenty-nine surface sediment samples from Effingham Inlet, a small fjord on the west coast of Vancouver Island, British Columbia, were analyzed for diatoms. This fjord has been selected for paleoceanographic investigation due to the presence of laminated sediments resulting from the dysoxic to anoxic bottom water conditions in the inner and outer basins of the inlet. Distributional patterns of the diatom microflora reflected proximity to littoral regions, phytoplankton production, and marine influence from outside the fjord. Principal components and cluster analyses of the microflora established four diatom assemblages with a clear separation between the inner and outer basin diatom floras. Inner basin stations were characterized by elevated absolute abundance with assemblages dominated by spring–early summer bloom taxa including Skeletonema costatum, Thalassiosira nordenskioeldii, and Thalassiosira pacifica. Chaetoceros spp. resting spores were abundant throughout Effingham Inlet, with the exception of the stations closest to the fjord head. Stations located in the outer basin and towards the fjord mouth had relatively lower absolute abundance yet showed a higher relative and absolute abundance of Thalassionema nitzschioides, Rhizosolenia setigera, Coscinodiscus radiatus, Ditylum brightwellii, Odontella longicruris, and Paralia sulcata in relation to the inner basin. Many of these latter taxa are often associated with late summer and autumn conditions in fjords along coastal British Columbia. Oceanographic data for Effingham Inlet suggest that increased offshore penetration is more likely to occur from summer to early fall, with a more restricted offshore influence in the inner basin. Diatom surface sediment assemblages in Effingham Inlet appear to reflect incursions of offshore waters into the fjord. Absolute abundance estimates and the preservation of lightly silicified taxa suggest excellent preservation of fossil material in the sediments of the predominantly anoxic inner basin. Preservation in the outer basin is reduced, reflecting more frequent recharge by oxygenated waters spilling over the outer sill into the basin. Our findings suggest the inner basin should be an optimal site for reconstruction of diatom production, with records from the outer basin providing more consistent information about offshore influence and coastal upwelling conditions over the Holocene. Estimates of diatom abundance within the inner basin sediment may serve as a good proxy of production, although proxy tracers of bottom water conditions and sedimentological analyses must be coupled to the diatom record to ensure depositional conditions were not influencing valve preservation or abundance. Our results suggest that fjords can serve as good environments for paleoceanographic reconstructions of both inshore and offshore conditions although careful site selection and understanding of processes affecting the microfossil record are essential.     

Calcareous nannofossil surface sediment assemblages from the Sicily Channel (central Mediterranean Sea): Palaeoceanographic implications

Quantitative analysis of 67 calcareous nannofossil assemblages from surface sediments recovered in a wide area across the Sicily Channel has been carried out in order to improve the interpretation of palaeontological data based on this planktonic group in a key area for Mediterranean palaeoceanographic studies. The investigation focused on three case studies that demonstrate the high potentiality of such a combined approach, taking into account the recent distribution of taxa or groups of taxa on the sea floor and the palaeontological record. The distribution of reworked specimens over the northern Sicily Channel sea floor validates the role of southern Sicily as a source region for reworked nannofossils and the role of rivers as their carrier. Eustatic sea-level fluctuations can be considered to be the main factor that influenced the abundance variations in sedimentary sequences of this area. The distribution of Florisphaera profunda can be explained in terms of topography (positive correlation) and mesoscale oceanographic circulation. In particular, its significant anti-correlation to the amount of chlorophyll-A deduced by satellite imagery validates the use of this species as a proxy for palaeoproductivity reconstructions. Finally, high abundance values of G. oceanica are confined to the westernmost part of the Sicily Channel, coinciding with a water mass salinity minimum. In particular, abundances of up to about 10% were observed in the westernmost part of the African Margin, suggesting the importance of the Atlantic Tunisian Current, whose activity is more pronounced in winter. The comparison of data of this species between 135 and 110 kyr BP, inside and outside the Channel, led us to deduce that the physical transport in almost unmodified waters of Atlantic origin might be the most important factor for its significant occurrence.     

Phylogenetic composition of bacterioplankton assemblages from the Arctic Ocean

We analyzed the phylogenetic composition of bacterioplankton assemblages in 11 Arctic Ocean samples collected over three seasons (winter-spring 1995, summer 1996, and summer-fall 1997) by sequencing cloned fragments of 16S rRNA genes. The sequencing effort was directed by denaturing gradient gel electrophoresis (DGGE) screening of samples and the clone libraries. Sequences of 88 clones fell into seven major lineages of the domain Bacteria: alpha(36%)-, gamma(32%)-, delta(14%)-, and epsilon(1%)-Proteobacteria; Cytophaga-Flexibacter-Bacteroides spp. (9%); Verrucomicrobium spp. (6%); and green nonsulfur bacteria (2%). A total of 34% of the cloned sequences (excluding clones in the SAR11 and Roseobacter groups) had sequence similarities that were <94% compared to previously reported sequences, indicating the presence of novel sequences. DGGE fingerprints of the selected samples showed that most of the bands were common to all samples in all three seasons. However, additional bands representing sequences related to Cytophaga and Polaribacter species were found in samples collected during the summer and fall. Of the clones in a library generated from one sample collected in spring of 1995, 50% were the same and were most closely affiliated (99% similarity) with Alteromonas macleodii, while 50% of the clones in another sample were most closely affiliated (90 to 96% similarity) with Oceanospirillum sp. The majority of the cloned sequences were most closely related to uncultured, environmental sequences. Prominent among these were members of the SAR11 group. Differences between mixed-layer and halocline samples were apparent in DGGE fingerprints and clone libraries. Sequences related to alpha-Proteobacteria (dominated by SAR11) were abundant (52%) in samples from the mixed layer, while sequences related to gamma-proteobacteria were more abundant (44%) in halocline samples. Two bands corresponding to sequences related to SAR307 (common in deep water) and the high-G+C gram-positive bacteria were characteristic of the halocline samples.     

Ice algal assemblages and vertical export of organic matter from sea ice in the Barents Sea and Nansen Basin (Arctic Ocean)

Algal communities and export of organic matter from sea ice were studied in the offshore marginal ice zone (MIZ) of the northern Barents Sea and Nansen Basin of the Arctic Ocean north of Svalbard by means of ice cores and short-term deployed sediment traps. The observations cover a total of ten stations within the drifting pack ice, visited over a period of 3 years during the period of ice melt in May and July. Maximum flux of particulate organic carbon and chlorophyll a from the ice at 1 m depth (1,537 mg C m⁻² per day and 20 mg Chl a m⁻² per day) exceeded the flux at 30 m by a factor of 2 during spring, a pattern that was reversed later in the season. Although diatoms dominated the ice-associated algal biomass, flagellates at times revealed similarly high biomass and typically dominated the exported algal carbon. Importance of flagellates to the vertical flux increased as melting progressed, whereas diatoms made the highest contribution during the early melting stage. High export of ice-derived organic matter and phytoplankton took place simultaneously in the offshore MIZ, likely as a consequence of ice drift dynamics and the mosaic structure of ice-covered and open water characteristic of this region.     

Assessment of surface sediment dynamics and response of benthic macrofauna assemblages in Boughrara Lagoon (SW Mediterranean Sea)

A study was conducted in the spring of 2009, the winters of 2010 and 2013, and in the summer of 2012 at 13 stations in Boughrara Lagoon, Tunisia (southern Mediterranean). The country⿿s largest lagoon, it is considered to be an anthropogenically stressed area, though a major tourist centre⿿Djerba Island⿿is located along its northern shores. The lagoon bottoms were studied via analyses of grain size, surface sediment composition, total organic matter (TOM) and through the trophic and functional organisation of benthic macrofauna. The results indicate that the bottoms are composed of fine, medium or coarse sands and that sediment distribution is controlled by water movement. Estimation of TOM content revealed that the studied samples present both normal and imbalanced sediments. The structure and organisation of the lagoon⿿s benthic macrofauna are dominated by select deposit feeders and underwent significant changes during the period 2010⿿2013.Subjected for decades to increased pollution due to growing human activities in the surrounding area, Boughrara Lagoon now appears to be impacted by certain environmental/anthropogenic stressors, as indicated by the presence of pollution-tolerant bio-indicator species in the imbalanced area. The response of the lagoon ecosystem to changes in benthic sediment deposition provides a potential assessment tool for similar habitats elsewhere.     

Phylogenetic composition of Arctic Ocean archaeal assemblages and comparison with Antarctic assemblages

Archaea assemblages from the Arctic Ocean and Antarctic waters were compared by PCR-denaturing gradient gel electrophoresis (DGGE) analysis of 16S rRNA genes amplified using the Archaea-specific primers 344f and 517r. Inspection of the DGGE fingerprints of 33 samples from the Arctic Ocean (from SCICEX submarine cruises in 1995, 1996, and 1997) and 7 Antarctic samples from Gerlache Strait and Dallman Bay revealed that the richness of Archaea assemblages was greater in samples from deep water than in those from the upper water column in both polar oceans. DGGE banding patterns suggested that most of the Archaea ribotypes were common to both the Arctic Ocean and the Antarctic Ocean. However, some of the Euryarchaeota ribotypes were unique to each system. Cluster analysis of DGGE fingerprints revealed no seasonal variation but supported depth-related differences in the composition of the Arctic Ocean Archaea assemblage. The phylogenetic composition of the Archaea assemblage was determined by cloning and then sequencing amplicons obtained from the Archaea-specific primers 21f and 958r. Sequences of 198 clones from nine samples covering three seasons and all depths grouped with marine group I Crenarchaeota (111 clones), marine group II Euryarchaeota (86 clones), and group IV Euryarchaeota (1 clone). A sequence obtained only from a DGGE band was similar to those of the marine group III Euryarchaeota:     

Regional patterns of nematode assemblages in the Arctic deep seas

Surface primary productivity and carbon flux in the Arctic Seas are higher along the warm Spitsbergen Water Current than along the ice-infested East Greenland Current. These contrasting oceanographic conditions are reflected in the deep-sea environment and may shape nematode assemblages. However, the paucity of samples in the Arctic deep seas precludes any regional scale assessment. In the present study, nematode assemblages were investigated in relation to a range of environmental variables along the 2,000 m isobath between latitudes 72°N and 79°N for both East and Western margins of the northern North Atlantic. Results showed that both margins had distinct environmental characteristics, with respect to chloroplastic pigments, sediment water content, sediment-bound organic matter, phospholipids and particulate proteins. Nematode assemblages varied according to these environmental changes. Along the more oligotrophic western margin, chloroplastic pigments increased towards the North, while the other environmental variables, nematode abundances and species richness decreased. In contrast, along the eastern margin, we observed higher quantities of organic matter and particulate protein, which supported higher abundance and species richness. Nematode assemblages along both margins varied according to food availability with species composition more variable in areas with lower amounts. Seventy percent of the species occurred in both margins indicating a low turnover of species. The present results support the hypothesis of a positive latitudinal gradient across the North Atlantic and further suggest that contemporary climate and recent ecological processes may predict nematode diversity patterns at larger scales.     

Seasonal and interannual dynamics of diatom assemblages in Sacrower See (NE Germany): a sediment trap study

Diatom assemblages from sediment trap samples collected during ten intervals between October 2003 and October 2005 in Sacrower See (NE Germany) were related to limnological and meteorological data. Sacrower See is a dimictic, 38 m deep, hypertrophic lowland lake (29.5 m a.s.l.). We identified distinct seasonal and interannual changes of diatom assemblages for the studied period. Diatoms showed a typical seasonal succession for temperate, dimictic, and eutrophic lakes. Stephanodiscus parvus, S. hantzschii, S. neoastraea, and S. alpinus had high accumulation rates during winter and spring, whereas species of the genera Stephanodiscus, Fragilaria, and Nitzschia were the predominant diatoms during summer and autumn. In a Canonical Correspondence Analysis, precipitation, air and water temperatures, epilimnetic calcium, pH, and total phosphorus concentrations together explained 70% of the variance of the diatom data. Interannual variability in the diatom assemblages during the two sampled years mainly seems to reflect changes in the total phosphorus concentration and temperature and secondarily the onset of the growing season and of stratification. Handling editor: J. Saros     

Sea Ice-Upper Ocean Ecosystems and Global Changes in the Arctic

In the Arctic, a clear temperature trend toward warming has been observed during the last two decades. The warming has led to a reduction in the area and thickness of the Arctic sea ice cover because of increased melting. Melt water is accumulated within the 0- to 30-m surface layer, markedly changing its hydrological and hydrochemical characteristics: this water is warmer and fresher, with lower nutrient concentrations. A stable thermocline at depths of 30–35 m weakens vertical mixing of water and hampers active exchange between nutrient-enriched water lying below the thermocline and nutrient-poor water under ice. This affects the qualitative and quantitative composition of upper-ocean phyto- and zooplankton. The changes in the climate and the ice environment cause marked alterations in the composition and structure of the biological communities of sea ice and under-ice surface water. Comparative analysis of materials collected in the anticyclonic Arctic Gyre at the ice stations Severnyi Polyus-22 (1975– 1981) and SHEBA (1997–1998) shows that over the last two decades the number of species of ice diatoms markedly decreased and the role of freshwater algae increased. The number of nematodes, copepods, amphipods, and turbellarians, which were the dominant groups in perennial ice, also declined sharply. The results suggest that the sea ice-upper ocean ecosystem is changing from typically marine to brackish-water. The main reason for these changes is probably global warming in the Arctic.     

Assessing the efficacy of chironomid and diatom assemblages in tracking eutrophication in High Arctic sewage ponds

Eutrophication is the most common water quality issue affecting freshwaters worldwide. Paleolimnological approaches have been used in temperate regions to track eutrophication over time, placing changes in historical context. Diatoms (Bacillariophyta) have a direct physiological response to changes in nutrients and are effective indicators of lake trophic status. Chironomids (Diptera) have also been used to track nutrient conditions; however, given that nutrients and oxygen are often tightly linked, it is difficult to disentangle which variable is driving shifts in assemblages. Here, we analyze chironomid and diatom remains in sediments from sewage-impacted ponds in the High Arctic. These ponds have the unusual characteristics of elevated nutrient and oxygen concentrations, unlike those of typical eutrophic lakes where deepwater oxygen is often depleted. Our data show that while diatom assemblages responded to changing nutrients, no concomitant changes in chironomid assemblage composition were recorded. Furthermore, the dominance of oligotrophic, cold stenothermic chironomid taxa, and lack of so-called “eutrophic” species in the eutrophic sewage ponds suggests that oxygen, not nutrients, structures chironomid assemblages at these sites.     

Composition and distribution of the pelagic and sympagic algal assemblages in the Laptev Sea during autumnal freeze-up

The phytoplankton and ice algal assemblages in the SiberianLaptev Sea during the autumnal freeze-up period of 1995 aredescribed. The spatial distribution of algal taxa (diatoms,dinoflagellates, chrysophytes, chlorophytes) in the newly formedice and waters at the surface and at 5 m depth differed considerablybetween regions. This was also true for algal biomass measuredby in situ fluorescence, chlorophyll (Chl) a and taxon-specificcarbon content. Highest in situ fluorescence and Chl a concentrations(ranging from 0.1 to 3.2 µg l^–1) occurred in surfacewaters with maxima in Buor Khaya Bay east of Lena Delta. Thealgal standing stock on the shelf consisted mainly of diatoms,dinoflagellates, chrysophytes and chlorophytes with a totalabundance (excluding unidentified flagellates <10 µm)in surface waters of 351–33 660 cells l^–1. Highestalgal abundance occurred close to the Lena Delta. Phytoplanktonbiomass (phytoplankton carbon; PPC) ranged from 0.1 to 5.3 µgC l^–1 in surface waters and from 0.3 to 2.1 µg Cl^–1 at 5 m depth, and followed the distribution patternof abundances. However, the distribution of Chl a differed considerablyfrom the distribution pattern shown by PPC. The algal assemblagein the sea ice, which could not be quantified due to high sedimentload, was dominated by diatom species, accompanied by dinoflagellates.Thus, already during the early stage of autumnal freeze-up,incorporation processes, selective enrichment and subsequentgrowth lead to differences between surface water and sea icealgal assemblages.     

Evaluating diatom community composition in the absence of marked limnological gradients in the high Arctic: a surface sediment calibration set from Cornwallis Island (Nunavut,Canada)

Fossil diatom assemblages preserved within the sedimentary record in Arctic lakes provide the potential to reconstruct past changes in important limnological variables. During the summers of 1992 and 1993, we examined previously unstudied freshwater ecosystems on Cornwallis Island, Arctic Canada, with the specific objectives of (1) documenting the limnology and modern diatom assemblages from this region, and (2) determining which environmental variables most influence diatom species distributions. The Cornwallis Island study sites displayed the least amount of variance in measured water chemistry variables in comparison to nearly all of our labs’ previous freshwater surveys in the Arctic. The small limnological gradients precluded the development of a statistically robust diatom inference model, but perhaps more importantly, allowed us to explore variations in diatom composition in the absence of marked variations in water chemistry. Diatom species turnover was minimal, with the most common diatom taxa being Achnanthidium minutissima, Nitzschia perminuta, N. frustulum, with lesser percent abundances of Chaemaepinnularia soehrensis, Navicula chiarae, Psammothidium marginulata, and A. kryophila. A small number of study sites differed from the majority with respect to water chemistry (e.g., coastal sites with high specific conductivities) and habitat availability (e.g., ephemeral ponds with extensive moss habitats), and these sites had markedly different diatom assemblages. These data reinforce previous observations that water chemistry and other climate-related factors are the primary environmental controls influencing diatom distributions at high latitudes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Distribution patterns of polychaete fauna in an Arctic fjord (Hornsund, Spitsbergen)

Polychaetes are one of the most important groups of benthic organisms in marine ecosystems. They dominate on the Arctic shelf and play an important role in ecosystem functioning. This study focuses on the polychaete biodiversity and their distribution patterns in Hornsund, an open glacial fjord, in western Spitsbergen and provides important baseline data for future studies of temporal fluctuations in benthic fauna. The main aim of this study was to assess how the polychaete abundance, biomass, diversity, community structure, and function vary along the Hornsund fjord’s axis, in relation to the environmental factors. Eighty-eight polychaete taxa were identified; an average density was 457 ind. m^?2 ± 237.5 SD. Three assemblages were distinguished (INNER, MIDDLE, and OUTER) along the fjord axis, reflecting the intensity of glacial disturbance. A clear division between the highly disturbed inner part of the fjord (Brepollen) and the less impacted middle and outer parts was observed. Continuous gradients in abundance, biomass, and diversity were found with all those values diminishing toward the inner region of the fjord. The polychaete assemblages' indices were significantly correlated with bottom temperature, sediment characteristics (grain size), and distance to the glacier (longitude). No significant correlations were found with depth or total organic carbon content. Carnivore and motile surface deposit feeding polychaete species dominated in the areas close to the glaciers, while the OUTER community was dominated by carnivores and surface sessile and discretely motile species, and had more complex trophic structure, with multiple species representing different functional groups including carnivores, sessile, discretely motile, and motile surface deposit feeders and motile burrowers.