Distinctive C isotopic signature distinguishes a novel sea ice biomarker in Arctic sediments and sediment traps

A C₂₅ highly branched isoprenoid (HBI) monoene hydrocarbon, designated IP₂₅, has been proposed previously to originate from diatoms living in Arctic sea ice, while the presence of IP₂₅ in sediments has been suggested to be a proxy for the occurrence of former Arctic sea ice. Here, we show that the ¹³C isotopic composition of IP₂₅ in sea ice, in sediment trap material collected under sea ice, and in high latitude northern sediments, is distinctive (isotopically ‘heavy’) and distinguishable from that of organic matter of planktonic or terrigenous origin. Mean δ¹³C values for IP₂₅ were − 22.3 ± 0.4‰ (sea ice), − 19.6 ± 1.1‰ (sediment traps) and − 19.3 ± 2.3‰ (sediments). These measurements, therefore, support further the proposed use of IP₂₅ as an Arctic sea ice proxy.     

闽江口沉积硅藻分布特征及其沉积意义

The distribution of diatoms in surface sediments in the Minjiang Estuary, southeast China, was investigated in2009. Total 56 species and other species belonging to 25 genera were identified, among them 11 species were dominant over 5%. Dominant species included Actinocyclus ehrenbergii, Coscinodiscus curvatulus, C. divisus, C.jonesianus, C. radiatus, C. rothii, C. subtilis, Cyclotella stylorum, Epithemia hyndmanii, Hydrosera whampoensis,and Trachyneis aspera. Diatom abundance varied spatially, with the absolute abundance of diatoms ranging from13 valves/g to 11×104 valves/g, and averaging 2.5×104 valves/g. A canonical correspondence analysis(CCA) was used to explain the relationships between diatom distribution and sediment properties in the Minjiang Estuary.CCA revealed that the major elements(Fe_2O_3, Na_2O, CaO, MgO, TiO_2, SiO_2, Al_2O_3, and K_2O) were closely related to diatom abundance. Four diatom assemblages were distinguished, representing different sediment properties,which may assist late Quaternary palaeoceanographic reconstructions of the Minjiang Estuary.     

Contribution of benthic and epiphytic diatoms to clam and oyster production in the Akkeshi-ko estuary

The species composition and dynamics of phytoplankton in the water column and its contribution to clam, Ruditapes philippinarum, and oyster, Crassostrea gigas, production were studied in the Akkeshi-ko estuary, eastern part of Hokkaido, Japan. A total 128 taxa of diatoms were identified, with 103 and 102 species occurring on the surface sediments and in the water column, respectively. Amphora sp., Bacillaria paradoxa var paxilifer, Cocconeis scutellum, Navicula sp., Nitzschia sigma, Paralia sulcata, Rhoicosphenia curvata, Synedra ulna and Thalassiosira sp. were most common and dominant in all stations in both water column and surface sediment. Benthic diatoms were most dominant in both water column and surface sediment. The species composition of epiphytes was, in part, similar to diatom assemblage of the water column and also to that of the surface sediment. The benthic and pelagic ratio in the diatom assemblage of the water column was very high. Benthic diatoms were able to resuspend into water column by turbulence stimulated by wind. Chlorophyll a concentration in the water column increased as wind speed increased. Seventy and 67% of the gut contents oysters consisted of benthic diatoms in 2003 and 2004 and also 78 and 87% of clams in 2003 and 2004, respectively. The availability of benthic and epiphytic diatom assemblages contributes significantly to food resources for clam and oyster mariculture due to resuspension by disturbance in the Akkeshi-ko estuary.     

Settling fluxes of diatoms to the interior of the Antarctic circumpolar current along 170 °W

An array of four sediment trap moorings recorded the particulate flux across the Antarctic Circumpolar Current (ACC) at 170 °W, between November 1996 and January 1998, as part of the US JGOFS-Antarctic Environment and Southern Ocean Process Study (AESOPS) program. The trap locations represent sampling within the Polar Frontal Zone, the Antarctic Polar Front, the Antarctic Zone and the Southern Antarctic Zone. Here we report observations from 1000 m below the sea-surface compared to seafloor and surface water distributions. Sub-sample splits from each trap were obtained and total diatom flux and species composition were determined. The diatom fluxes were quantified using both a dilution and a ‘spike’ method to allow for the rapid repeatability of measurements. Diatom flux was found to be highly seasonal across the ACC particularly at higher latitudes. Marine snow aggregates of intact diatom cells and chains were the major components of the biogenic flux. Siliceous particle size was noted to decrease with increasing latitude, which could be aligned with a shift of the diatom assemblage to small-size species/sea-ice affiliated species. A ‘double-structured’ diatom flux was recorded at the location of the Antarctic Polar Front trap, with a shift in the diatom assemblage from larger to smaller diatoms in the second flux episode. The sediment trap assemblage shows deviations from the surface water assemblage, while surface sediment samples indicate that significant dissolution occurs after 1000 m and at the sediment–water interface. Estimation of diatom biovolumes across the ACC shows that large diatoms have the potential to greatly impact biogenic fluxes to the ocean interior despite their low fluxes. Small species of the genus Fragilariopsis could potentially export as much C_org as Fragilariopsis kerguelensis near the retreating ice edge. However, their low abundance in the surface sediments also suggests that these diatoms are a shallow export species.     

广西钦州湾外湾表层沉积硅藻分布特征

本文对钦州湾外湾54个表层沉积物中硅藻进行分析,鉴定出硅藻153种,分属50属。其中Thalassionema nitzschioides是本区域内最为丰富的硅藻种类。对应分析结果表明,钦州湾表层沉积硅藻分布主要受到海水盐度的影响,其中淡水硅藻Achnanthes delicatula、A.hauckiana、A.levanderi和Cocconeis disculus可以作为低盐度海水环境的指示种,半咸水种Cyclotella striata/C.stylorum、Paralia sulcata与海洋中硅藻Thalassionema frauenfeldii、T.nitzschioides则可以很好的指示高盐度环境。钦州湾外湾54个表层沉积站位可划分为3个硅藻分布区,分别对应高低不同的海水盐度,与实际站位地理分布基本一致。     

The widespread occurrence of highly branched acyclic C20, C25 and C30 hydrocarbons in recent sediments and biota—A review

The occurrence of widely distributed acyclic isoprenoid hydrocarbons (C_2O, C₂₅ and C₃₀) with highly branched structures, in sediments and biota is reviewed. The compounds occur as alkanes and alkenes with from one to at least four double bonds in young aquatic (both marine and lacustrine) sediments from many parts of the globe (e.g. Peru, Antarctica, Gulf of Suez, North Sea, Atlantic) sometimes in high concentrations (e.g. 40 μg g⁻¹ sediment) relative to other hydrocarbons. However, the compounds rapidly disappear in older sediments, possibly due to biodegradation and reaction with sedimentary sulphur. These and other factors are discussed.The sources of this fascinating group of hydrocarbons remain largely unknown, though evidence points to algae (possibly diatoms) as one possibility. The recent identification of related sulphur-containing compounds in a crude oil promises to extend the number of reports of these compounds still further, and to increase their importance as environmental biological markers.     

Distribution and germination of viable diatom resting stage cells in sediments of the East China Sea

The abundance and temporal patterns of viable diatom resting stage cells in sediments of the East China Sea in 2006 have been investigated. The abundance of viable resting stages was enumerated with the most probable number (MPN) technique. Overall, 25 diatom species (including varieties) belonging to 16 genera were detected. Viable resting stage cells were common, on the order of 10³ to 10⁶ cells/g dry mass. The abundant taxa included Skeletonema marina, S. dohrnii, Chaetoceros curvisetus and Thalassiosira spp., with Skeletonema accounting for 83.3% of the cell. The effects of temperature (10, 15, 20, 25℃), salinity (20, 25, 30, 35 psu) and light intensity (0, 300, 4 000, 8 000 lux) on the germination of resting stages in sediment samples were also determined. The results showed that light intensity had a significant positive effect on the germination of diatom resting stage cells (p <0.01) and that diatoms could not germinate in darkness. Temperature and salinity had no noticeable effects on the germination. The present study suggests that there are abundant diatom resting cells in sediments of the East China Sea, which could serve as a "seed bank" in the phytoplankton population succession and contribute to the initiation of algal blooms.     

Temporal and spatial variability in export production in the SE Pacific Ocean: evidence from siliceous plankton fluxes and surface sediment assemblages

Flux of siliceous plankton and taxonomic composition of diatom and silicoflagellate assemblages were determined from sediment trap samples collected in coastal upwelling-influenced waters off northern Chile (30°S, CH site) under “normal” or non-El Niño (1993–94) and El Niño conditions (1997–98). In addition, concentration of biogenic opal and siliceous plankton, and diatom and silicoflagellate assemblages preserved in surface sediments are provided for a wide area between 27° and 43°S off Chile. Regardless of the year, winter upwelling determines the maximum production pattern of siliceous microorganisms, with diatoms numerically dominating the biogenic opal flux. During the El Niño year the export is markedly lower: on an annual basis, total mass flux diminished by 60%, and diatom and silicoflagellate export by 75%. Major components of the diatom flora maintain much of their regular seasonal cycle of flux maxima and minima during both sampling periods. Neritic resting spores (RS) of Chaetoceros dominate the diatom flux, mirroring the influence of coastal-upwelled waters at the CH trap site. Occurrence of pelagic diatoms species Fragilariopsis doliolus, members of the Rhizosoleniaceae, Azpeitia spp. and Nitzschia interruptestriata, secondary components of the assemblage, reflects the intermingling of warmer waters of the Subtropical Gyre. Dictyocha messanensis dominates the silicoflagellate association almost year-around, but Distephanus pulchra delivers ca. 60% of its annual production in less than three weeks during the winter peak. The siliceous thanatocoenosis is largely dominated by diatoms, whose assemblage shows significant qualitative and quantitative variations from north to south. Between 27° and 35°S, the dominance of RS Chaetoceros, Thalassionema nitzschioides var. nitzschioides and Skeletonema costatum reflects strong export production associated with occurrence of coastal upwelling. Both highest biogenic opal content and diatom concentration at 35° and 41°–43°S coincide with highest pigment concentrations along the Chilean coast. Predominance of the diatom species Thalassiosira pacifica and T. poro-irregulata, and higher relative contribution of the silicoflagellate Distephanus speculum at 41°–43°S suggest the influence of more nutrient-rich waters and low sea surface temperatures, probably associated with the Antarctic Circumpolar Water.     

我国典型潮间带底栖硅藻群落空间分布特征

底栖硅藻是河口、泥质海滩生态系统的重要生物类群,其数量与分布变化直接或间接地影响到多种海洋生物的饵料供给、海鸟的生存与迁徙。通过对我国从南到北14个典型潮间带表层沉积物中底栖硅藻的分析,共鉴定出硅藻153种,其物种多样性（Shannon-Weaver 指数范围:1.61~4.39）与生物量（范围:0.09×103 ~ 10×103个/g,干质量）在空间上存在显著差异;聚类分析与相似性分析检验（r=0.738,P=0.1%< 0.01）表明,14个调查区域的硅藻群落在空间上可分为4个大的组合,分别对应着渤、黄、东、南海4个区域,吻合了我国近海浮游植物的地理分布特征。与环境因素的相关性分析表明,底栖硅藻群落的物种多样性与生物量受沉积物类型与盐度的影响显著,泥质区域更有利于硅藻的保存,而生物多样性在泥砂质低盐区域相对较高。此外,底栖硅藻生物量与表层沉积物中TOC含量表现出显著正相关性;δ13C数值的空间变化范围为-25‰~-20‰,指示了有机质陆海混合的特征,其中,涧河河口、黄河口、大沽河口、苏北浅滩区域,δ13C数值明显偏正,表明底栖硅藻可能是这些区域沉积物中有机碳的重要贡献者。     

Biogeochemistry of sterols in plankton, settling particles and recent sediments in a cold ocean ecosystem (Trinity Bay, Newfoundland)

In the context of a multidisciplinary study to determine current and past ecosystem health and the relative contributions of sources of organic matter (marine vs. terrestrial and natural vs. anthropogenic input), sterols were determined in plankton, settling particles and sediments from Trinity Bay, Newfoundland, a sub-polar Atlantic Ocean ecosystem. The centric diatoms Chaetoceros spp., Thalassiosira spp. and Leptocylindrus danicus were all prominent in the plankton samples, and centric diatoms predominated in the settling particles. Plankton samples contained 0.4±0.4 mg/g dw (1995) or 1.4±1.3 mg/g dw (1996) total sterols, with cholesta-5,24-dien-3β-ol (mean 26% of total sterols), cholest-5-en-3β-ol (24%) and cholesta-5,22(E)-dien-3β-ol (13%) chief among these, denoting diatom and zooplankton sources. In settling particles, the prominence of cholesta-5,24-dien-3β-ol (24%), cholest-5-en-3β-ol (24%), cholesta-5,22(E)-dien-3β-ol (13%) and 24-methylcholesta-5,22(E)-dien-3β-ol (9%) again suggested mainly marine sources. The sterol composition of plankton and settling particles from different sampling periods showed a high degree of consistency. Higher plant C₂₉ sterols (notably 24-ethylcholest-5-en-3β-ol, 9–26%) were prominent in sediments from both inshore and offshore sites. No decreasing trend in total or individual sterols was observed down the 30-cm sediment cores, suggesting good overall preservation. No 5β-stanols such as 5β-cholestan-3β-ol (coprostanol) were detected in offshore sediments, plankton or settling particles, with only low levels (5β-cholestan-3β-ol max. 4.4%, 5β-cholestan-3α-ol max. 5.1%) in certain inshore sediments. This suggests that raw sewage discharges in rural Newfoundland are being efficiently degraded or dispersed, or that inputs are highly localized. Source apportionation of organic matter in the sediment samples based on sterol composition was attempted. This highlighted the large terrestrial contribution to the sterols in marine sediments (up to 58% of sterols inshore, 24% offshore) and suggests either degradation or effective recycling of marine sterols.     

Experiments on the resuspension of estuarine sediments containing benthic diatoms

A laboratory system was used to test the effect of water flow on the resuspension of mud and sand sediments and, specifically, benthic diatoms from the Ems estuary, The Netherlands. Current velocities generated by two rotating cylinders in a cylindrical tank were determined by a small float and a laser Doppler velocimeter. At low angular velocities, the amount of suspended matter increased linearly with angular velocity and the float current velocity. However, at higher angular velocities, the increase in current velocity was less because of the strong turbulence: concomitantly, the current velocity boundary layer (δ) became thinner and the suspended matter concentration increased rapidly. The dominant diatom species from the sandy sediment were suspended in two distinct groups, one of which consisted of the species Navicula aequorea, Navicula salinicola, Ophephora martyi and Opephora pacifica, and was more exclusively bound to sand grains than the other. The benthic diatom species inhabiting the silty sediment did not show this difference. The most important shortcoming in the experiments was the inability to determine the radial and vertical velocity components. This precluded reliable calculations of the shear stress. The data presented emphasize the importance of finding a method to determine the shear stress under experiments and field conditions so that direct comparisons can be made. Despite this it is assumed that, just as under the experimental conditions discussed, under natural conditions in shallow waters resuspension starts at current velocities as low as ca. 10 cm s⁻¹.     

Observations of resuspended diatoms in the turbid tidal edge

Observations of resuspended diatoms in the shallow waters (<60 cm) of the turbid tidal edge are described for single sites on two tidal flats–the Molenplaat in the Westerschelde estuary, and the Hond in the Ems-Dollard estuary, The Netherlands. High concentrations of chlorophyll-a (chl-a) were observed in the trailing edge of the ebbing tide in water depths of <20 cm, after which concentrations decreased markedly. Peak mean values were 19 μg chl-a l⁻¹ in 10 cm of water at the Molenplaat, and 45 μg chl-a l⁻¹ in 5 cm of water at the Hond. Similar trends were observed on the flooding tide, although peak values were far less pronounced (6 and 30 μg chl-a l⁻¹ respectively). Microscopic examination of the diatom community within the turbid tidal edge at the Molenplaat revealed that peaks in biomass were caused by suspended benthic diatoms, as well as the large centric diatom Coscinodiscus sp., particularly on the ebb tide. Planktonic diatoms other than Coscinodiscus sp. were more randomly distributed and did not appear to follow any particular trend. It would seem that as the tide recedes, resuspended benthic diatoms and large Coscinodiscus sp. cells become concentrated in the shallow water. However, the virtual absence of Coscinodiscus sp. from the leading edge of the flooding tide suggests that most of the resuspended cells do not settle to the seabed, but are washed away into the channels. The small peak of benthic diatoms at the leading edge of the flood tide is most likely resuspended locally from the sediment, along with large numbers of diatom frustules.     

Sources and transport of lipids on Amazon continental shelf

Dissolved material and recent sediment from the Amazon continental shelf have been analyzed for hydrocarbons to study the sources and potential fate of the transported organic matter. Dissolvedn-alkanes are present at low concentrations (ppb level) and are dominated by lipids from marine phytoplankton with carbon number maxima (C_max) at C₁₈/C₂₂ and an even-to-odd carbon predominance < C₃₀ (CPI_17–27 from 0.18 to 0.54). In the sediments, bimodal distributions ofn-alkane chain length suggest a mixed input of terrestrial (C_max at C₂₇/C₂₉/C₃₁ and CPI_25–33 from 0.75 to 1.82) and phytoplanktonic/microbial (C_max at C₂₀ and CPI_15–25 from 0.38 to 0.62) organic matter. Sesquiterpenes were the most significant cyclic compounds in all the dissolved samples analyzed reflecting a contribution from resinous trees to the terrestrial organic pool. On the other hand, enhanced concentrations of these compounds in the dissolved phase on the northwest portion of the Amazon shelf, contrasting with decreased concentrations in the sediment samples, suggest that dissolved lipids are released from solid phase in the intensely stirred seabed. Structured organic matter in the sediment has been characterized as being composed of, on average: 19% plant cuticles, 25% woody tissue, 13% pollens and spores, 24% amorphous material, 7% bituminite and 12% altered organic material.     

Sources and preservation of organic matter in Plum Island salt marsh sediments (MA, USA): long-chain n-alkanes and stable carbon isotope compositions

Elemental (TOC, TN, C/N) and stable carbon isotopic (δ¹³C) compositions and n-alkane (nC_16–38) concentrations were measured for Spartina alterniflora, a C₄ marsh grass, Typha latifolia, a C₃ marsh grass, and three sediment cores collected from middle and upper estuarine sites from the Plum Island salt marshes. Our results indicated that the organic matter preserved in the sediments was highly affected by the marsh plants that dominated the sampling sites. δ¹³C values of organic matter preserved in the upper fresh water site sediment were more negative (−23.0±0.3‰) as affected by the C₃ plants than the values of organic matter preserved in the sediments of middle (−18.9±0.8‰) and mud flat sites (−19.4±0.1‰) as influenced mainly by the C₄ marsh plants. The distribution of n-alkanes measured in all sediments showed similar patterns as those determined in the marsh grasses S. alterniflora and T. latifolia, and nC₂₁ to nC₃₃ long-chain n-alkanes were the major compounds determined in all sediment samples. The strong odd-to-even carbon numbered n-alkane predominance was found in all three sediments and nC₂₉ was the most abundant homologue in all samples measured. Both δ¹³C compositions of organic matter and n-alkane distributions in these sediments indicate that the marsh plants could contribute significant amount of organic matter preserved in Plum Island salt marsh sediments. This suggests that salt marshes play an important role in the cycling of nutrients and organic carbon in the estuary and adjacent coastal waters.     

东海表层沉积硅藻

1977年国家海洋局组织所属单位对东海大陆架进行了综合调查,采集了比较丰富的样品.有关资料报导了在东海大陆架有12种硅藻的记录,大陆坡11种,海槽区12种,如归并重复出现的种类,总计有20种硅藻.本文基本上用同一批材料对沉积硅藻的沉积问题,提出了我们的一些看法.     

THE VERTICAL MIGRATION OF DIATOM IN LITTORAL OF XIBIAN AND XIAO''AO

The vertical migration of diatoms in mud-flats was studied in littoral of Xibian, Longhai County and Xiao'ao, Lianjiang County, the dominant species being Gyrosigma spencerii and Hantzschia virgata respectively. Both light and tide are important factors affecting the vertical migration of diatoms. With the advent of tide, the diatom community disappears from the surface. It reappears gradually when exposed to light. In the dark, the diatoms never appear in great quantity on the surface in laboratory or in littoral. Whether in the light or in the dark, the diatoms aggregate in maximum numbers in the upper 2 mm of the sediment. But when the tide has covered the sediments for 4 hr, Hantz. virgata penetrates the mud down to a depth of 14 mm, Gy. spencerii, 12 mm. We have found that Gy. spencerii posseses phototaxis, but no tide rhythm and no day-night rhythm in our laboratory.     

The effect of suspended sediment concentration on the settling velocity of cohesive sediment in quiescent water

A laboratory study was carried to qualitatively investigate the effect of suspended sediment concentration C on the settling velocity w_s of cohesive sediment in quiescent water. A bay mud sample was mixed with water in a cylindrical container, and three optical back scatterance sensors were then used to measure suspended sediment concentrations of the mud–water mixture at three levels every 15 s for 5 h while sediments were settling in the quiescent water. Based on the measured sediment concentrations, the settling velocities at different concentrations were derived from the depth-integrated mass balance equation. This study has found that the settling velocity w_s is independent of C in the free settling regime of C<0.3 g/l, and then increases nonlinearly with C in the enhanced settling regime of 0.3<C<4.3 g/l, and finally decreases sharply with C in the hindered settling regime of C>4.3 g/l. The maximum settling velocity occurs at C≈4.3 g/l and is about nine times faster than the settling velocity in the free settling regime. A single empirical formula is also proposed to calculate the settling velocities at different sediment concentrations.     

冲绳海槽晚第四纪硅藻生物地层划分与对比

本文对冲绳海槽中段B孔岩芯中的沉积硅藻进行了分析。共鉴定出52属166种和变种．根据硅藻属种组成、热带远洋种的相对含量以及整个硅藻植物群的变化特征共划分出9个组合带，并对它们的特征进行了描述；结合^14C测年数据。对该岩芯的地层年代归属进行了确定，全新世与晚更新世的界线位于该孔125cm处．上部地层为全新统，下部地层为上更新统；同时与邻近DGKS9603钻孔的硅藻生物地层进行了对比．     

Higher plant n-alkane average chain length as an indicator of petrogenic hydrocarbon contamination in marine sediments

The n-alkane average chain length (ACL) is the weight-averaged number of carbon atoms of the higher plant C₂₅–C₃₃ n-alkanes. The abundance of individual n-alkanes from higher plant sources generally increases with increasing carbon number in coastal marine sediments around Taiwan, but this trend is reversed for petrogenic hydrocarbons. The ACL would potentially be lowered if petrogenic hydrocarbons were added to sediments containing biogenic hydrocarbons alone. To test this idea, a marine environment off southwestern Taiwan known to contain both biogenic and petrogenic hydrocarbons and two nearby rivers were selected for investigating possible difference in ACL values between their sediments. The average CPI of C₂₅–C₃₃ n-alkanes was 4.08 ± 2.04 (range 1.90–8.96, n = 15) for the river sediments and 1.70 ± 0.16 (range 1.43–1.97, n = 15) for the marine sediments. The ACL of C₂₅–C₃₃ n-alkanes for river sediments ranged from 29.2 to 30.5 (average 29.9 ± 0.4), and for marine sediments from 28.4 to 29.3 (average 28.9 ± 0.3). The ACL difference between marine and river sediments was significant (Student's t test at 99% confidence) although it appeared small. It is suggested that the ACL can be an additional indicator for detection of petrogenic hydrocarbons in coastal marine sediments.     

Fossil diatom assemblages as paleoecological indicators of paleo-water environmental change in the Ulleung Basin,East Sea,Republic of Korea

The fossil diatom assemblage record from two sediment cores obtained from the Ulleung Basin, East Sea, Republic of Korea, revealed changes in the diatom assemblage zones in PG1 and PD3 core samples. The two sediment cores were δC¹⁴ dated and approximately represented the late Pleistocene–Holocene. The analysis of age zones in the PG1 core and PD3 core was assessed based on the frequency of variations, and occurrences of biostratigraphical fossil diatom species. During the Last Glacial Maximum (LGM), the sea level was lower than that at present and the Ulleung Basin became isolated from the Pacific Ocean. As a result, there would have been a limited Tsushima Warm Current (TWC) influence, and salinity would have decreased resulting in increased freshwater and coastal diatoms. The distribution pattern of diatoms presented in the cores was associated with changes in water temperature and salinity and the adding of terrigenous material brought about by the input of freshwater. Changes in the abundance of a tychopelagic diatom, Paralia sulcata, reflected the effect of the water currents. Diatom temperature (Td) values and the ratio of centric/pennate diatoms provided evidence of limited influences of the TWC and freshwater inflow. It is thought that all assemblage zones were influenced by the TWC, which had an important effect on the distribution and composition of fossil diatoms.