Seasonal growth and senescence in continental shelf ecosystems: a test of the SEEP hypothesis

The hypothesis that continental shelf ecosystems export a major fraction of the carbon produced by the phytoplankton during the spring bloom was tested during the Shelf Edge Exchange Processes (SEEP) experiment off the north-east coast of the United States in 1984. The total benthic standing stocks in terms of organic carbon (macrofauna, meiofauna and bacteria) have been estimated in the SEEP area. Their preponderance on the continental shelf was partial evidence that little organic matter escapes to the upper continental slope. Measurements of the metabolism of the biota allowed calculation of turnover times of organic detritus and the total biota. The turnover time of detritus increased as grain size decreased, suggesting that fine-grained deposits contain mostly refractory, non-reactive compounds, especially on the deep slope. Turnover times of the total biota were about the same in the coarse- as in the fine-grained shelf deposits, but a far larger fraction of the turnover was attributed to the bacteria in the fine sediments. On average, about 25 per cent of the primary production appeared to be utilized by the aerobic benthos on the continental shelf in the SEEP area. The role of anaerobes at depth in the sediments remains uncertain. This study, along with a review of other recent work, leads to the conclusion that only a small fraction of continental shelf phytodetritus is exported across a distinct shelf-slope hydrographic frontal system. What is not consumed in the spring is utilized on the shelf during the ensuing stratified season. More open ended ecosystems may export production more readily.     

Factors regulating benthic food chains in tropical river deltas and adjacent shelf areas

Benthic food chains of the Amazon (Brazil) and Fly (Papua New Guinea) river deltas and adjacent shelves are compared. Abundance patterns of the major trophic groups (bacteria, meiofauna, and macroinfauna) are similar between regions, with very low densities, or the absence of benthos, within and near the deltas. For muds in the more quiescent areas, benthic abundance and productivity are highest, commonly coinciding with maximum pelagic primary production. Episodes of physical disturbance, erratic food supply, and dilution of river-derived, particulate organic matter foster the development of opportunistic benthic communities of variable diversity and low biomass, dominated by bacteria. These pioneering assemblages are the main food of penaeid shrimp, which dominate the demersal trawl fisheries of both fluvial-dominated regions.     

Metazoan meiofauna biomass, grazing, and weight-dependent respiration in the Northern Gulf of Mexico deep sea

Metazoan meiofauna are ubiquitous in marine soft sediments and play a pivotal role in diagenesis of particulate organic matter. However, the relative importance of meiofauna to the function of deep-sea benthic boundary layer communities has not been resolved. Here, meiofauna biomass, respiration, and grazing on aerobic heterotrophic bacteria were estimated and compared to standing stocks and fluxes of other benthic components (e.g., bacteria and macrofauna). Biomass and respiration declined with depth. Highest biomass and respiration occurred in the proximity of the Mississippi River on the upper continental slope of the central Gulf of Mexico. Meiofauna required 7% of their biomass per day to meet their metabolic energy budget, compared to approximately 24% day⁻¹ in shallow water. Respiration accounted for 8–22% of whole sediment community respiration (SCOC), reflecting the importance of meiofauna in diagenesis, deep-sea carbon budgets, and global biogeochemical cycles.     

Potential effects of temperature on the benthic infaunal community on the southeastern Bering Sea shelf: Possible impacts of climate change

In the late 1950s, Soviet researchers collected benthic infaunal samples from the southeastern Bering Sea shelf. Approximately 17 years later, researchers at University of Alaska Fairbanks also sampled the region to assess infaunal biomass and abundance. Here, the two data sets were examined to document patterns and reveal any consistent differences in infaunal biomass among major feeding groups between the two time periods. No significant differences in the geometric mean biomass of all taxa pooled were indicated between the two study periods (1958–1959=49.1 g m⁻²; 1975–1976=60.8 g m⁻²; P=0.14); however, significant differences were observed for specific functional groups, namely carnivores, omnivores and surface detritivores. Of the 64 families identified from both data sets from all functional groups, 21 showed statistically significant (P0.05) differences in mean biomass. Of the 21 families showing significant differences, 19 (91%) of the families had higher mean biomass in the 1975–1976 data set. The above differences suggest a trend toward higher overall infaunal biomass for specific functional groups during mid 1970s compared with the late 1950s. Temperature measurements and literature data indicate that the mid-1970s was an unusually cold period relative to the period before and after, suggesting a mechanistic link between temperature changes and infaunal biomass. Food-web relationships and ecosystem dynamics in the southeastern Bering Sea indicate that during cold periods, infaunal biomass will be elevated relative to warm periods due to elevated carbon flux to the benthos and exclusion of benthic predators on infaunal invertebrates by the cold bottom water on the shelf. As long-term observations of temperature and sea-ice cover indicate a secular warming trend on the Bering Sea shelf, the potential changes in food-web relationships could markedly alter trophic structure and energy flow to apex consumers, potentially impacting the commercial, tourist and subsistence economies.     

Vertical Distribution of Meiobenthos in Bathyal Sediments of the Eastern Mediterranean Sea: Relationship with Labile Organic Matter and Bacterial Biomasses

Abstract. Quantitative information on the vertical distribution of meiofaunal abundances and biomass were obtained from samples collected at 3 bathyal stations in the Eastern Mediterranean Sea located at the same depth but characterized by different food supply. Vertical distribution patterns of nieiofauna were investigated in relation to the biochemical composition of the sediment organic matter (proteins, carbohydrates, and lipids) and compared to benthic bacterial standing stocks. No significant relationship between bacteria and meiofauna was found, whereas a significant relationship between protein and lipid concentrations and total meiofauna density was observed. These data suggest that labile organic matter. considered as material readily aVdihbk to benthic consumers, may be an important factor regulating meiofaunal abundance and vertical distribution in deep-sea sediments.     

Infaunal benthic communities of the New Zealand continental shelf

Seventeen infaunal benthic communities are recognised from sampling on the New Zealand continental shelf and upper continental slope. These are characterised in terms of geographic distribution, substrate and bathymetric range, and commoner infaunal bivalves and echinoderms.     

The importance of bioturbation to continental slope sediment structure and benthic processes off Cape Hatteras, North Carolina

Even though the continental slope off Cape Hatteras has sediment accumulation rates on the order of 1 cm/year, large areas of soft sediment are intensively reworked by infaunal organisms. Primary sedimentary structures have been completely replaced with biogenic structures. Surface sedimentary structures are dominated by the bioturbational activities of a deep burrowing infauna (to at least 30 cm). The layer actively mixed by the benthos, as evidenced by sediment profile and X-ray images, is estimated to range from 5 to 20 cm, with the residence time for particles within the surface mixed layer ranging from about 4.5 to 18 years. The biological mixing parameter (G) ranges from 0.4 to 5.5, which indicates moderate to strong biological mixing relative to accumulation and strata formation. Bioturbation contributes to the dynamic forces affecting the surface sediments by decreasing compaction of sediment layers and dilating sediment fabrics by sediment mixing, and introducing large water-filled burrows and voids to subsurface sediments. The sediment profile images captured numerous subsurface feeding voids, and worms in the process of making deep burrows, many of which extended below the 5 to 10 cm average depth of the apparent color redoxpotential discontinuity layer. High rates of accumulation of organic-rich sediment lead to high standing stocks of benthos and intensive feeding/burrowing activity that result in organic rich stratagraphic sequences that are thoroughly mixed. Cape Hatteras is a apparent focusing point for the transport of shallow water sediments to the deep sea. Sediments across other areas of the continental slope just 100's of kilometers south of Cape Hatteras are not as thoroughly mixed or biologically active.     

Surficial sediments of the wave-dominated Orange River Delta and the adjacent continental margin off south-western Africa

The textural and compositional characteristics of the surficial shelf sediments north and south of the Orange River Delta are reviewed and compared. Sediments are fractionated and dispersed both north- and southwards of the Orange River mouth by wave action, longshore drift and subsurface currents. The mean grain size decreases both offshore and southwards in response to decreased wave influence at the seabed and the competence of the weak poleward undercurrent respectively. The increasing dominance of marine biogenic components in sediments south of the prodelta indicates a greater marine influence, modifying previous inferences that the Namaqualand mudbelt is primarily derived from the southward transport of Orange River sediments. A sharp distinction can be drawn between sediments of the Orange Shelf to the south and the Walvis Shelf to the north. Foraminiferarich deposits that dominate the Orange middle shelf and slope indicate that upwelling is an inner-shelf phenomenon. On the Walvis Shelf, foraminiferal sediments are confined to the slope and outer shelf. Fish debris is more common in Walvis Shelf sediments. Although phosphorite and glauconite sands often occur together in the same deposits on the Orange Shelf, the two minerals are concentrated in separate deposits on the Walvis Shelf.     

Preliminary study on community structures of meiofauna in the middle and eastern Chukchi Sea

Sediment core samples were collected from 17 stations in the middle and eastern Chukchi Sea during the sixth Chinese National Arctic Research Expedition(CHINARE-Arctic) in summer 2014.The samples were analyzed for composition,abundance,biomass,vertical distribution,size spectra,and ecological indexes of meiofauna.A total of 14 meiofauna taxa were detected,and the free-living marine nematodes comprised the most dominant taxon,accounting for 97.21% of the average abundance.The abundance and biomass of meiofauna were within ranges of(218.12±85.83)-(7 239.38±1 557.15) ind./(10 cm~2) and(130.28±52.17)-(3 309.56±1 751.80) μg/(10 cm~2),with average values of(2 391.90±1 966.19) ind./(10 cm~2) and(1 549.73±2 042.85) μg/(10 cm~2)(according to dry weight)respectively.Furthermore,91.26% of the individuals were distributed in the top layer of 0-5 cm of surface sediment,and 90.84% had sizes of 32-250 μm.Group diversity index of meiofauna in the survey area was low,and the variation of abundance was the main difference in meiofauna communities among all stations.Abundance and biomass of meiofauna were not significantly correlated with environmental factors except concentration of nutrient Si in bottom seawater.Abundance of meiofauna in shallow water of marginal seas in the Pacific sector of the Arctic Ocean is likely at a same level and higher than that in most of China sea areas,suggesting that the shallow water of the summer Chukchi Sea is a continental shelf area with rich resources of meiofauna.The Chukchi Sea is important for studying the ecosystem of the Arctic Ocean and environmental responses.However,studies on meiofauna in the Chukchi Sea are still not enough,and in the future,natural and human disturbances may increase due to global warming,the Arctic channel opening,and other factors.Thus,more studies on meiofauna should be required,in order to know more about how the Arctic benthic community would alter.     

辽河口邻近海域小型底栖生物的空间分布及季节变化

本文研究了辽河口邻近海域2013年8月、10月和2014年5月3个航次小型底栖生物的种类及其空间分布,分析了小型底栖生物丰度和生物量的季节变化。结果表明,3个航次（夏季、秋季和春季）小型底栖生物的平均丰度分别为（264±83） ind/（10 cm2）、（216±85） ind/（10 cm2）和（227±67） ind/（10 cm2）,平均生物量分别为（272±125）μg/（10 cm2）、（207±89）μg/（10 cm2）和（244±103）μg/（10 cm2）。与其他研究海域相比,辽河口小型底栖的丰度和生物量处于较低水平。共鉴定出了14个小型生物类群,按照丰度排序,线虫是最优势的类群,夏季、秋季和春季3个航次占总丰度的比例分别为94.0%、92.5%和90.8%;其他优势类群为多毛类、桡足类和双壳类。小型底栖生物量的优势类群则为多毛类（41.1%~44.0%）,高于线虫（33.8%~36.5%）,其次是双壳类（2.6%~6.7%）。水平分布的研究表明,调查海域近岸入海口小型底栖生物的丰度和生物量普遍低于近海海域,但是秋季时近岸分布与近海差距不大。垂直分布的研究表明,95.9%的小型底栖生物分布于0~5 cm的表层沉积物中。小型底栖生物的丰度和生物量在夏季时都达到高峰值。与环境因子的相关分析表明,小型底栖生物的数量分布与盐度和水深呈极显著正相关（P<0.01）,与叶绿素a呈显著正相关（P<0.05）。     

Comparative biomass structure and estimated carbon flow in food webs in the deep Gulf of Mexico

A budget of the standing stocks and cycling of organic carbon associated with the sea floor has been generated for seven sites across a 3-km depth gradient in the NE Gulf of Mexico, based on a series of reports by co-authors on specific biotic groups or processes. The standing stocks measured at each site were bacteria, Foraminifera, metazoan meiofauna, macrofauna, invertebrate megafauna, and demersal fishes. Sediment community oxygen consumption (SCOC) by the sediment-dwelling organisms was measured at each site using a remotely deployed benthic lander, profiles of oxygen concentration in the sediment pore water of recovered cores and ship-board core incubations. The long-term incorporation and burial of organic carbon into the sediments has been estimated using profiles of a combination of stable and radiocarbon isotopes. The total stock estimates, carbon burial, and the SCOC allowed estimates of living and detrital carbon residence time within the sediments, illustrating that the total biota turns over on time scales of months on the upper continental slope but this is extended to years on the abyssal plain at 3.6 km depth. The detrital carbon turnover is many times longer, however, over the same depths. A composite carbon budget illustrates that total carbon biomass and associated fluxes declined precipitously with increasing depth. Imbalances in the carbon budgets suggest that organic detritus is exported from the upper continental slope to greater depths offshore.The respiration of each individual “size” or functional group within the community has been estimated from allometric models, supplemented by direct measurements in the laboratory. The respiration and standing stocks were incorporated into budgets of carbon flow through and between the different size groups in hypothetical food webs. The decline in stocks and respiration with depth were more abrupt in the larger forms (fishes and megafauna), resulting in an increase in the relative predominance of smaller sizes (bacteria and meiofauna) at depth. Rates and stocks in the deep northern GoM appeared to be comparable to other continental margins where similar comparisons have been made.     

Abundance and biomass of meiobenthos in the spawning ground of anchovy (Engraulis japanicus) in the southern Huanghai Sea

1 Introduction Meiofauna is an important group in benthic small food web energetically due to their small size, high abundance and fast turnover rates. The production of meiofauna is equal to or higher than that of macrofau- na in estuaries, shallow waters and deep sea (Gerlach, 1971; Platt and Warwick, 1980; Heip et al., 1985; Zhang et al., 2004). A role of meiofauna may be the recycling of nutrients. Marine nematodes may keep the bacterial colonies on sand grains in active phase of growth …     

Infaunal macrobenthos of the oxygen minimum zone on the Indian western continental shelf

The Arabian Sea is characterized by a mid‐depth layer of reduced dissolved oxygen (DO) concentration or oxygen minimum zone (OMZ ‐DO concentration <0.5 ml·l^?1) at ~150–1000 m depth. This OMZ results from the flux of labile organic matter coupled with limited intermediate depth water ventilation. Generally, benthic animals in the OMZ have morphological and physiological adaptations that maximize oxygen uptake in the limited oxygen availability. Characteristics of OMZ benthos have been described from only a few localities in the Arabian Sea. We measured the bottom water DO and studied the characteristics of infaunal macrobenthos of the Indian western continental shelf by collecting samples at 50, 100 and 200 m in depth from 7° to 22° N. The DO values observed at 200 m (0.0005–0.24 ml·l^?1) indicated that this area is lying within an OMZ. Five major taxa, namely Platyhelminthes, Sipunculoidea, Echiuroidea, Echinodermata and Cephalochordata were absent from the samples collected from this OMZ. In general, declines in total macrobenthic density and biomass and polychaete species richness and diversity were observed in this OMZ compared with the shallower depths above it. Community analyses of polychaetes revealed the dominance of species belonging to families Spionidae, Cirratulidae and Paraonidae in this OMZ. Low oxygen condition was more pronounced in the northern continental shelf edge (≤0.03 ml·l^?1), where the majority of spionids including Prionospio pinnata and cirratulids were absent; whereas amphipod, isopod and bivalve communities were not impacted.     

Small benthic size classes along the N.W. European Continental Margin: spatial and temporal variability in activity and biomass

In the context of the European OMEX Programme this investigation focused on gradients in the biomass and activity of the small benthic size spectrum along a transect across the Goban Spur from the outer Celtic Sea into Porcupine Abyssal Plain. The effects of food pulses (seasonal, episodic) on this part of the benthic size spectrum were investigated. Sediments sampled during eight expeditions at different seasons covering a range from 200 m to 4800 m water depth were assayed with biochemical bulk measurements: determinations of chloroplastic pigment equivalents (CPE), the sum of chlorophyll a and its breakdown products, provide information concerning the input of phytodetrital matter to the seafloor; phospholipids were analyzed to estimate the total biomass of small benthic organisms (including bacteria, fungi, flagellata, protozoa and small metazoan meiofauna). A new term ‘small size class biomass' (SSCB) is introduced for the biomass of the smallest size classes of sediment-inhabiting organisms; the reduction of fluorescein-di-acetate (FDA) was determined to evaluate the potential activity of ester-cleaving bacterial exoenzymes in the sediment samples.At all stations benthic biomass was predominantly composed of the small size spectrum (90% on the shelf; 97–98% in the bathyal and abyssal parts of the transect). Small size class biomass (integrated over a 10 cm sediment column) ranged from 8 g C m⁻² on the shelf to 2.1 g C m⁻² on the adjacent Porcupine Abyssal Plain, exponentially decreasing with increasing water depth. However, a correlation between water depth and SSCB, macrofauna biomass as well as metazoan meiofauna biomass exhibited a significantly flatter slope for the small size classes in comparison to the larger organisms.CPE values indicated a pronounced seasonal cycle on the shelf and upper slope with twin peaks of phytodetrital deposition in mid spring and late summer. The deeper stations seem to receive a single annual flux maximum in late summer. SSCB and heterotrophic activity are significantly correlated to the amount of sediment-bound pigments. Seasonality in pigment concentrations is clearly followed by SSCB and activity. In contrast to macro- and megafauna which integrate over larger periods (months/years), the small benthic size classes, namely bacteria and foraminifera, proved to be the most reactive potential of the benthic communities to any perturbations on short time scales (days/weeks). The small size classes, therefore, occupy a key role in early diagenetic processes.     

南大西洋洋中脊调查区小型底栖生物丰度和生物量研究

“大洋一号”调查船于2011年5—6月在南大西洋中脊14°S附近进行了7个站位的小型底栖生物采样。共鉴定出小型底栖生物10个类群。小型底栖生物平均丰度为(60.63±54.77) ind/10 cm²,平均干重生物量为(9.42±8.92) μg/10 cm²。线虫是其中的优势类群,丰度为(47.42±47.99)ind/10 cm²,占总丰度的78.21%,另外,肉鞭动物和桡足类分别占总丰度的16.63%和3.91%。生物量前3位的类群依次为桡足类、线虫和肉鞭动物。小型底栖生物密度随沉积物深度增加而减少,约73.55%的生物丰度分布在0～2 cm层内。个体大小方面,有75.32%的小型底栖生物粒径处于32～125 μm范围内。     

渤海底栖桡足类群落结构的研究

渤海1998年9月航次的调查结果.小型底栖生物总平均丰度为(8688±5097)个/m2,底栖桡足类居第2位,平均丰度为(663±569)个/m2,占总数量的76%.根据77种底栖桡足类丰度所做的聚类和标序分析将渤海20个站位划分为4个组合.根据11种环境因子数据所做的聚类和主成分分析将研究海域划分为两个生境、4个亚区.对研究海域进行的分区与自然分区是基本一致的.支配研究海域底栖桡足类群落结构的主要环境因子是水深和沉积物粒度.     

Dense infaunal assemblages on the continental slope off Cape Hatteras, North Carolina

Unusually dense assemblages of benthic infaunal invertebrates have been discovered in continental slope sediments off Cape Hatteras, North Carolina. Densities were highest on the upper slope, ranging from 24,055 to 61,244 (X¯=46,255) individuals m⁻² in nine samples taken at a 600-m site in 1984 and 1985, and from 15,522 to 89,566 (X¯=37,282) individuals m⁻² in single samples at 15 stations over a wider depth range of 530 to 1535 m in 1992. A lower slope station at 2000 m sampled six times in 1984–1985 and again in 1992, had densities consistently higher than 8500 individuals m⁻². Species richness and diversity are consistently lower on the Cape Hatteras slope than at other locations off North Carolina and elsewhere in the western North Atlantic. The 1992 studies indicated that the upper slope infaunal assemblages (600m) were dominated by oligochaetes, while the middle slope assemblages (800–1400 m) were dominated by the polychaeteScalibregma inflatum. This latter depth range could be defined into two assemblages based upon suites of less abundant species. At depths of 1500–2000 m, a lower slope assemblage dominated by various deposit feeding polychaetes and oligochaetes was found. Results from the 1984–1985 studies suggest seasonal or year-to-year patterns in the dominance ofS. inflatum andCossura longocirrata. Unusually high sedimentation rates and organic carbon flux have been recorded from the slope off Cape Hatteras and may account for the high infaunal productivity in the area. Most of the dominant infaunal organisms are species more typical of shallow, coastal habitats rather than deep-sea species that dominate other areas of the U.S. Atlantic continental slope. Parallel investigations regarding the nature of organic matter in the Cape Hatteras sediments have revealed a mixture of both marine and terrestrially derived carbon, only a small percentage of which is composed of the smaller molecular weight polyunsaturated fatty acids more typical of continental slope sediments. It is likely that the high percentage of refractory organic matter would favor the survival of preadapted shelf species over those from adjacent slope environments.     

Oxygenation episodes on the continental shelf of central Peru: Remote forcing and benthic ecosystem response

The interplay between the oxygen minimum zone and remotely-forced oxygenation episodes determines the fate of the benthic subsystem off the Central Peruvian coast. We analyzed a 12 year monthly time-series of oceanographic and benthic parameters at 94 m depth off Callao, Central Peru (12°S), to analyze: (i) near-bottom oxygen level on the continental shelf in relation to dynamic height on the equator (095°W); and (ii) benthic ecosystem responses to oxygen change (macrobiotic infauna, meiofauna, and sulphide-oxidizing bacteria, Thioploca spp.). Shelf oxygenation episodes occurred after equatorial dynamic height increases one month before, consistent with the propagation of coastal trapped waves. Several but not all of these episodes occurred during El Niños. The benthic biota responded to oxygenation episodes by undergoing succession through three major ecological states. Under strong oxygen deficiency or anoxia, the sediments were nearly defaunated of macro-invertebrates and Thioploca was scarce, such that nematode biomass dominated the macro- and meiobiotas. When frequency of oxygenation events reduced the periods of anoxia, but the prevailing oxygen range was 10–20 μmol L⁻¹, mats of Thioploca formed and dominated the biomass. Finally, with frequent and intense (>40 μmol L⁻¹) oxygenation, the sediments were colonized by macrofauna, which then dominated biomass. The Thioploca state evolved during the 2002–2003 weak EN, while the macrofauna state was developed during the onset of the strong1997–1998 EN. Repeated episodes of strong oxygen deficiency during the summer of 2004, in parallel with the occurrence of red tides in surface waters, resulted in the collapse of Thioploca mats and development of the Nematode state. Ecological interactions may affect persistence or the transition between benthic ecosystem states.     

Multi-decadal synthesis of benthic–pelagic coupling in the western arctic: Role of cross-shelf advective processes

Using geographic information systems (GIS) software and geostatistical techniques, we utilized three decades of water-column chlorophyll a data to examine the relative importance of autochthonous versus allochthonous sources of reduced carbon to benthic communities that occur from the northern Bering to the eastern Beaufort Sea shelf. Spatial trend analyses revealed areas of high benthic biomass (>300 g m⁻²) and chlorophyll (>150 mg m⁻²) on both the southern and northern Chukchi shelf; both areas are known as depositional centers for reduced organic matter that originates on the Bering Sea shelf and is advected northward in Anadyr and Bering shelf water masses. We found a significant correlation between biomass and chlorophyll a in the Chukchi Sea, reflective of the strong benthic–pelagic coupling in a system that is utilized heavily by benthic-feeding marine mammals. In contrast, there was no significant correlation between biomass and chlorophyll in the Beaufort Sea, which by comparison, is considerably less productive (biomass and chlorophyll, <75 g m⁻² and <50 mg m⁻², respectively). One notable exception is an area of relatively high biomass (50–100 g m⁻²) and chlorophyll (80 mg m⁻²) near Barter Island in the eastern Beaufort Sea. Compared to other adjacent areas in the Beaufort Sea, the chlorophyll values in the vicinity of Barter Island were considerably higher and likely reflect a long-hypothesized upwelling in that area and close coupling between the benthos and autochthonous production. In the Bering Sea, a drop in benthic biomass in 1994 compared with previous measurements (1974–1993) may support earlier observations that document a decline in biomass that began between the 1980s and 1990s in the Chirikov Basin and south of St. Lawrence Island. The results of this study indicate that the benthos is an excellent long-term indicator of both local and physical advective processes. In addition, this work provides further evidence that secondary production on arctic shelves can be significantly augmented by reduced carbon advected from highly productive adjacent shelves.