Tectonic geomorphology of the Ryukyu Trench-Arc-Backarc System: geological-geophysical exploration and mapping

OKINAWA TROUGH; BASIN

Characterization of proteolytic bacteria from the Aleutian deep-sea and their proteases

Six deep-sea proteolytic bacteria taken from Aleutian margin sediments were screened; one of them produced a cold-adapted neutral halophilic protease. These bacteria belong to Pseudoalteromonas spp., which were identified by the 16S rDNA sequence. Of the six proteases produced, two were neutral cold-adapted proteases that showed their optimal activity at pH 7-8 and at temperature close to 35 degrees C, and the other four were alkaline proteases that showed their optimal activity at pH 9 and at temperature of 40-45 degrees C. The neutral cold-adapted protease E1 showed its optimal activity at a sodium chloride concentration of 2 M, whereas the activity of the other five proteases decreased at elevated sodium chloride concentrations. Protease E1 was purified to electrophoretic homogeneity and its molecular mass was 34 kDa, as estimated by sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE). The molecular weight of protease E1 was determined to be 32,411 Da by mass spectrometric analysis. Phenylmethyl sulfonylfluoride (PMSF) did not inhibit the activity of this protease, whereas it was partially inhibited by ethylenediaminetetra-acetic acid sodium salt (EDTA-Na). De novo amino acid sequencing proved protease E1 to be a novel protein.

Diversity and Distribution of Sediment NirS-Encoding Bacterial Assemblages in Response to Environmental Gradients in the Eutrophied Jiaozhou Bay, China

A gene-clone-library-based molecular approach was used to study the nirS-encoding bacteria-environment relationship in the sediments of the eutrophic Jiaozhou Bay. Diverse nirS sequences were recovered and most of them were related to the marine cluster I group, ubiquitous in estuarine, coastal, and marine environments. Some NirS sequences were unique to the Jiaozhou Bay, such as the marine subcluster VIIg sequences. Most of the Jiaozhou Bay NirS sequences had their closest matches originally detected in estuarine and marine sediments, especially from the Chesapeake Bay, indicating similarity of the denitrifying bacterial communities in similar coastal environments in spite of geographical distance. Multivariate statistical analyses indicated that the spatial distribution of the nirS-encoding bacterial assemblages is highly correlated with environmental factors, such as sediment silt content, NH4+ concentration, and OrgC/OrgN. The nirS-encoding bacterial assemblages in the most hypernutrified stations could be easily distinguished from that of the least eutrophic station. For the first time, the sedimentological condition was found to influence the structure and distribution of the sediment denitrifying bacterial community.

Tectonic Evolution and Dynamics of Deepwater Area of Pearl River Mouth Basin, Northern South China Sea

Quantitative studies on the evolution and dynamics of the deepwater area of Pearl River Mouth basin (PRMB) were carried out based on the latest geological and seismic data. The study area is generally in an extensional state during the Cenozoic. The major extension happened in the earlier syn-rift stages before 23 Ma and the extension after 23 Ma is negligible. Two rapid subsidence periods, 32-23 Ma and 5.3-2.6 Ma, are identified, which are related to the abrupt heat decay during margin breakup and the collision between the Philippine Sea plate and the Eurasian plate, respectively. The strongest crustal thinning in the Baiyun (sic) sag may trigger the syn-rift volcanism along the weak faulted belt around the sag. The Cenozoic tectonic evolution of the study area could be divided into five stages: rifting (similar to 50-40 Ma), rift-drift transition (similar to 40-32 Ma), early post-breakup (similar to 32-23 Ma), thermal subsidence (similar to 23-5.3 Ma) and neotectonic movement (similar to 5.3-0 Ma).

Genetic model of the hydrate system in the fine grain sediments in the northern continental slope of South China Sea

Gas hydrate samples were obtained firstly in China by drilling on the northern margin of South China Sea (SCS). To understand the formation mechanism of this unique accumulation system, this paper discusses the factors controlling the formation of the system by accurate geophysical interpretation and geological analysis, based on the high precision 2-D and 3-D multichannel seismic data in the drilling area. There are three key factors controlling the accumulation of the gas hydrate system in fine grain sediment: (1) large volume of fluid bearing methane gas Joins the formation of gas hydrate. Active fluid flow in the northern South China Sea makes both thermal gas and/or biogenic gas migrate into shallow strata and form hydrate in the gas hydrate stability zone (GHSZ). The fluid flow includes mud diapir and gas chimney structure. They are commonly characterized by positive topographic relief, acoustic turbidity and push-down, and low reflection intensity on seismic profiles. The gas chimneys can reach to GHSZ, which favors the development of BSRs. It means that the active fluid flow has a close relationship with the formation and accumulation of gas hydrate. (2) The episodic process of fracture plays an important role in the generation of gas hydrate. It may provide the passage along which thermogenic or biogenic gas migrated into gas hydrate stability zone (GHSZ) upward. And it increases the pore space for the growth of hydrate crystal. (3) Submarine landslide induced the anomalous overpressure activity and development of fracture in the GHSZ. The formation model of high concentration gas hydrate in the drilling sea area was proposed on the basis of above analysis.

Petroleum System in Deepwater Basins of the Northern South China Sea

The northern South China Sea margin has experienced a rifting stage and a post-rifting stage during the Cenozoic. In the rifting stage, the margin received lacustrine and shallow marine facies sediments. In the post-rifting thermal subsidence, the margin accumulated shallow marine facies and hemipelagic deposits, and the deepwater basins formed. Petroleum systems of deepwater setting have been imaged from seismic data and drill wells. Two kinds of source rocks including Paleogene lacustrine black shale and Oligocene-Early Miocene mudstone were developed in the deepwater basin of the South China Sea. The deepwater reservoirs are characterized by the deep sea channel rill, mass flow complexes and drowned reef carbonate platform. Profitable capping rocks on the top are mudstones with huge thickness in the post-rifting stage. Meanwhile, the faults developed during the rifting stage provide a migration path favourable for the formation of reservoirs. The analysis of seismic and drilling data suggests that the joint structural and stratigraphic traps could form giant hydrocarbon fields and hydrocarbon reservoirs including syn-rifting graben subaqueous delta, deepwater submarine fan sandstone and reef carbonate reservoirs.

Seismic characteristics of a reef carbonate reservoir and implications for hydrocarbon exploration in deepwater of the Qiongdongnan Basin, northern South China Sea

Our analysis of approximately 40,000 km of multichannel 2-D seismic data, reef oil-field seismic data, and data from several boreholes led to the identification of two areas of reef carbonate reservoirs in deepwater areas (water depth >= 500 in) of the Qiongdongnan Basin (QDNB), northern South China Sea. High-resolution sequence stratigraphic analysis revealed that the transgressive and highstand system tracts of the mid-Miocene Meishan Formation in the Beijiao and Ledong-Lingshui Depressions developed reef carbonates. The seismic features of the reef carbonates in these two areas include chaotic bedding, intermittent internal reflections, chaotic or blank reflections, mounded reflections, and apparent amplitude anomalies, similar to the seismic characteristics of the LH11-1 reef reservoir in the Dongsha Uplift and Island Reef of the Salawati Basin, Indonesia, which house large oil fields. The impedance values of reefs in the Beijiao and Ledong-Lingshui Depressions are 8000-9000 g/cc x m/s. Impedance sections reveal that the impedance of the LH11-1 reef reservoir in the northern South China Sea is 800010000 g/cc x m/s, whereas that of pure limestone in BD23-1-1 is > 10000 g/cc x m/s. The mid-Miocene paleogeography of the Beijiao Depression was dominated by offshore and neritic environments, with only part of the southern Beijiao uplift emergent at that time. The input of terrigenous sediments was relatively minor in this area, meaning that terrigenous source areas were insignificant in terms of the Beijiao Depression: reef carbonates were probably widely distributed throughout the depression, as with the Ledong-Lingshui Depression. The combined geological and geophysical data indicate that shelf margin atolls were well developed in the Beijiao Depression, as in the Ledong-Lingshui Depression where small-scale patch or pinnacle reefs developed. These reef carbonates are promising reservoirs, representing important targets for deepwater hydrocarbon exploration. (C) 2008 Elsevier Ltd. All rights reserved.

Gas hydrate occurrence on the continental slope of the northern South China Sea

High-resolution multi-channel seismic data and geological samples were collected during two research cruises of the R/V FENDOU 4 in 1999 and 2000. Studies on these data and samples together with results from sites 1143-1145 and 1148 of ODP Leg 184 suggest that the geological structure on the continental slope of the northern South China Sea is favorable for the formation of gas hydrates. Bottom simulating reflectors (BSRs) and geochemical anomalies which indicate the existence of gas hydrates have been recognized in sediments of the Xisha Trough, the Dongsha Rise and the accretionary wedge of the Manila subduction zone. These gas hydrates are generated by two different mechanisms depending on the tectonic regime and the seismic and geochemical characteristics. The first applies to the passive continental margin of the nor-them South China Sea on the Dongsha Rise and in the Xisha Trough. The gas hydrates are associated with diapiric structures, active faults, slumps and gravity flows as well as high Late Cenozoic sedimentation rates. Their seismic expression includes BSRs, seismic blanking zones and velocity anomalies. The second mechanism is operative on the active continental margin along the Manila subduction zone, especially in the accretionary wedge. Here, gas hydrate occurrence is marked by widespread BSRs and acoustic 'pull-down' possibly related to the existence of free gas in the sediments beneath the BSR. The thickness of the seismic blanking zones averages 250 m, suggesting that the stable gas hydrate zone has about the same thickness. (c) 2005 Elsevier Ltd. All rights reserved.

Geochemical properties of the variolitic andesite from Susong in the Dabie orogenic belt, China

The variolitic andesite from the Susong County in the Dabie Mountains implies that it was erupted in water. The mineralogy of the varioles is primarily radiate plagioclase (albite sind oligoclase), with little pyroxene, hornblende and quartz (derived from alteration). The pyroxene, hornblende and quartz are in the interstices between plagiocalse. The matrix consists of glass, hornblende, chlorite, epidote and zoisite. It is clearly subjected an extensive alteration. The andesite has an uncommon chemical composition. The SiO2 content is about 56.8%, TiO2 = 0.9%, MgO = 6.4%, Fe2O3 (tot) = 6.7%similar to 7.6%, 100Mg/(Mg + Fe) = 64.1 similar to 66.2. Mg-# is significantly high. The andesite has high abundances of large-lithophile trace elements (e.g. K, Ba. Sr, LREE), e.g. La/Nb = 5.56 similar to 6.07, low abundances of high-strength-field elements (HFSE e.g. Ta, Nb, P, Ti), particularly Ta and Nb strongly depleted. These are consistent with the characteristics of subduction-related magmas. In the spider diagram of trace elements, from Ce to right hand, the abundances of elements decrease quickly, showing a character of the continental margins. There has a strong punishment of light-rare-earth elements, with a significant diffraction of REEs (the mean value of (La/Yb)(N) is 32.84). No Eu anomaly, but there are anomaly high (La/Yb)(N) = 28.63 similar to 36.74, (La/Y)(N) = 70.33 similar to 82.4. The elements Y and Yb are depleted greatly, Y<20 g/g, Y-N = 2.74 similar to 2.84, Yb-N = 2.18 similar to 2.35. From the La-(La/Sm) diagram, the andesite is derived from partial melting. But the epsilone value of Nd is - 18.7 similar to -19.2, so that the material source may be the mantle materials affected by the crustal materials. The Nd model age is 1.9 Ga indicating that the variolitic basaltic andesite was resulted from the mantle wedge of North China block, which had the Nd model age of 2.5Ga, when the Yangze block which had the Nd model age of 1.7Ga subducted beneath it. So the variolitic andesite has characteristics of the island-are volconic rocks oil a continental basement in the vicinity of the destructive continental margin.

Major element, trace element, and Sr, Nd and Pb isotope studies of Cenozoic basalts from the South China Sea

The whole rock K-Ar ages of basalts from the South China Sea basin vary from 3.8 to 7.9 Ma, which suggest that intra-plate volcanism after the cessation of spreading of the South China Sea (SCS) is comparable to that in adjacent regions around the SCS, i.e., Leiqiong Peninsula, northern margin of the SCS, Indochina block, and so on. Based on detailed petrographic studies, we selected many fresh basaltic rocks and measured their major element, trace element, and Sr-Nd-Pb isotope compositions. Geochemical characteristics of major element and trace element show that these basaltic rocks belong to alkali basalt magma series, and are similar to OIB-type basalt. The extent of partial melting of mantle rock in source region is very low, and magma may experience crystallization differentiation and cumulation during the ascent to or storing in the high-level magma chamber. Sr-Nd-Pb isotopic data of these basaltic rocks imply an inhomogeneous mantle below the South China Sea. The nature of magma origin has a two end-member mixing model, one is EM2 (Enriched Mantle 2) which may be originated from mantle plume, the other is DMM (Depleted MORB Mantle). Pb isotopic characteristics show the Dupal anomaly in the South China Sea, and combined with newly found Dupal anomaly at Gakkel ridge in Arctic Ocean, this implies that Dupal anomaly is not only limited to South Hemisphere. In variation diagrams among Sr, Nd and Pb, the origin nature of mantle below the SCS is similar to those below Leiqiong peninsula, northern margin of the SCS and Indochina peninsula, and is different from those below north and northeast China. This study provides geochemical constraints on Hainan mantle plume.

Estimation of gas hydrate saturation using constrained sparse spike inversion: Case study from the northern South China Sea

Bottom-simulating reflectors (BSRs) were observed beneath the seafloor in the northern continental margin of the South China Sea (SCS). Acoustic impedance profile was derived by Constrained Sparse Spike Inversion (CSSI) method to provide information on rock properties and to estimate gas hydrate or free gas saturations in the sediments where BSRs are present. In general, gas hydrate-bearing sediments have positive impedance anomalies and free gas-bearing sediments have negative impedance anomalies. Based on well log data and Archie's equation, gas hydrate saturation can be estimated. But in regions where well log data is not available, a quantitative estimate of gas hydrate or free gas saturation is inferred by fitting the theoretical acoustic impedance to sediment impedance obtained by CSSI. Our study suggests that gas hydrate saturation in the Taixinan Basin is about 10 - 20% of the pore space, with the highest value of 50%, and free gas saturation below BSR is about 2 - 3% of the pore space, that can rise to 8 - 10% at a topographic high. The free gas is non-continuous and has low content in the southeastern slope of the Dongsha Islands. Moreover, BSR in the northern continental margin of the SCS is related to the presence of free gas. BSR is strong where free gas occurs.

Rifting process and formation mechanisms of syn-rift stage prolongation in the deepwater basin, northern South China Sea

Based on the latest seismic and geological data, tectonic subsidence of three seismic lines in the deepwater area of Pearl River Mouth Basin (PRMB), the northern South China Sea (SCS), is calculated. The result shows that the rifting process of study area is different from the typical passive continental margin basin. Although the seafloor spreading of SCS initiated at 32 Ma, the tectonic subsidence rate does not decrease but increases instead, and then decreases at about 23 Ma, which indicates that the rifting continued after the onset of seafloor spreading until about 23 Ma. The formation thickness exhibits the same phenomenon, that is the syn-rift stage prolonged and the post-rift thermal subsidence delayed. The formation mechanisms are supposed to be three: (1) the lithospheric rigidity of the northern SCS is weak and its ductility is relatively strong, which delayed the strain relaxation resulting from the seafloor spreading; (2) the differential layered independent extension of the lithosphere may be one reason for the delay of post-rift stage; and (3) the southward transition of SCS spreading ridge during 24 to 21 Ma and the corresponding acceleration of seafloor spreading rate then triggered the initiation of large-scale thermal subsidence in the study area at about 23 Ma.

热带西太平洋边缘晚第四纪以来的古环境研究

利用取自热带太平洋边缘的5个柱状岩芯Ph05-5、WP3、MD2908、Z14-6和WP7，通过对其微体古生物和生物地球化学成分的分析，结合浮游有孔虫氧、碳同位素数据、AMS14C测年结果和已有的研究资料，分别研究了热带西太平洋边缘晚第四纪以来的钙质超微化石群落特征、碳酸盐旋回、超微化石氧碳同位素特征、古生产力和上层海水结构演化以及高分辨率东亚夏季风指标，并探讨了碳酸盐旋回、古生产力和上层海水结构演化、全新世东亚夏季风强弱变化的控制机制，以及钙质超微化石下透光带属种Florisphaera profunda的相对百分含量变化在热带太平洋边缘的古环境指示意义及不同环境条件下的控制机制。 西菲律宾海Ph05-5岩芯和台湾东南部WP3岩芯中CaCO3和钙质超微化石碳酸盐溶解指数、初级生产力指标的变化显示约190 kyr BP以来CaCO3含量整体上都表现为冰期高、间冰期低的“太平洋型”旋回特征，但菲律宾以东海区在末次冰期（MIS 4~2期）内部又显示出间冰段含量高、冰段含量低的“大西洋型”旋回特征。碳酸盐旋回的主要控制因素在菲律宾以东海区是碳酸盐溶解作用，而台湾东南海区则是初级生产力变化引起的钙质生物输入量的波动。 西菲律宾海Ph05-5岩芯中190 kyr BP以来的钙质超微化石δ18O值在末次间冰期（MIS 5e）和全新世明显低于末次冰期（MIS 5d~2）和倒数第二次冰期（MIS 6）。超微化石δ18O值与浮游和底栖有孔虫δ18O值都呈明显的正相关关系，但是超微化石的δ18O平均值重于表层浮游有孔虫Globigeriniodes rubber，轻于中层Neogloboquadrina dutertrei。超微化石δ13C值变化与其绝对丰度变化表明西菲律宾海区大约在MIS 6和MIS 5e期，表层海水初级生产力相当稳定且显著低于其它各时期；约从MIS 5d期开始表层初级生产力显著上升，并一直持续到末次冰期；在MIS1、2期，表层初级生产力有所下降，但仍高于MIS 6和MIS 5e期。 Ph05-5岩芯中钙质超微化石下透光带属种F. profunda相对百分含量和浮游有孔虫转换函数的温跃层深度变化表明西太平洋暖池北缘约190 kyr BP 以来，营养跃层和温跃层冰期（MIS 6期和5d~2期）浅，间冰期（MIS5e）和全新世深，MIS5e期是最近两个冰期旋回中营养跃层和温跃层深度最深的时期。F. profunda百分含量初级生产力转换方程计算结果和与钙质超微化石绝对丰度的变化显示冰期初级生产力高，间冰期和全新世初级生产力低， MIS5e期初级生产力最低。此外，Δδ13C C. wullerstorf-coccolith和Δδ13C C. wullerstorf-N. dutertrei差值的变化也说明190 kyr BP以来冰期表层输出生产力高于间冰期，其中最为突出的特征也是MIS5e期为生产力输出的最低值期。上层海水结构和生产力的上述变化特征与现代La Niňa事件爆发时的海洋环境相当类似，该海区MIS5e期可能是La Niňa事件频繁爆发的一段时期，可以与现代La Niňa现象类比。同时，根据Wyrtki提出的信风张弛理论推测MIS5e期，由于La Niňa事件的频繁爆发黑潮主流应该是增强的。 热带西太平洋暖池核心区WP7岩芯中250 kyr BP以来的初级生产力在间冰期明显低于冰期，间冰期生产力相对稳定，冰期(MIS6，4和2)生产力升降幅度较大；冰期向间冰期过渡的冰消期内生产力的降低是相对缓慢的，而间冰期向冰期过渡时生产力的增加是迅速的。初级生产力指标的交叉频谱分析显示出偏心率、斜率、岁差（半岁差）、30kyr和50kyr周期。说明暖池区晚第四纪以来古生产力变动除了受全球冰量变动控制以外，低纬区的岁差驱动也是主要控制因子。约250 kyr BP以来古生产力的变化与Vostok冰心的CO2记录在30kyr周期上高度相关，低CO2值与高生产力相对应，印证了大洋初级生产力在全球碳循环中扮演了重要的“吸收者”的角色，30kyr周期是赤道太平洋地区生物生产力ENSO式变化特征的体现，这也证明了低纬地区生物泵在控制大气CO2浓度方面的重要作用。另外，在岁差周期段，底栖有孔虫U+B含量和N. dutertrei δ13C代表的生产力变动领先于有孔虫δ18O代表的冰体积变化约2.6~7kyr（0.5kyr,2.6 kyr,4.2 kyr, 7kyr），表明在千年时间尺度上日射是该区生产力变化的直接驱动力，两极冰体积变化（δ18O）没有明显的影响低纬气候波动。 对热带西太平洋边缘海区WP7、Ph05-5、WP3和Z14-6四个岩芯中晚第四纪以来超微化石下透光带属种F. profunda相对百分含量变化进行横向对比研究表明，F. profunda百分含量分布并非简单受水深控制，而是由海水上层复杂的生态因素所决定；F. profunda相对百分含量变化曲线与浮游有孔虫N. dutertrei氧同位素曲线呈不同程度的相关性，N. dutertrei 氧同位素越轻，对应的F. profunda相对百分含量越高；冲绳海槽中部的Z14-6岩芯中F. profunda相对含量变化的主控因子是黑潮流的强弱变化；WP7和Ph05-5两个岩芯中F. profunda相对含量变化与低纬地区暖池变化特征密切相关。 6800 yr BP以来台湾东北部MD2908岩芯中F. profunda相对百分含量变化与太阳黑子数量呈明显的反相关关系，而且3700 yrBP以来更为明显。由于F. profunda相对百分含量变化与该地区东亚夏季风引起的降雨量密切相关，所以推测6800 yr BP以来东亚夏季风受太阳活动影响，其中的纽带是IZCT的纬向位置变动。

南海北部陆缘深水区构造演化及其资源效应

南海北部陆缘深水区（水深>300m）蕴藏着丰富的资源，我国对深水区的地质研究刚刚起步，但相关领域已成为科研热点。深水油气盆地的构造演化是油气勘探中最重要的基础性研究之一，因此针对我国南海北部陆缘深水区开展构造演化及其资源效应的研究具有重要的理论意义和实际意义。 本文利用钻井和地震资料并结合区域地质资料，重点研究了珠江口盆地深水区的结构和构造演化，取得如下创新性成果：1）首次利用半地堑分析方法系统解剖了研究区的结构、各构造单元发育特征，在此基础上指出五个有利油气运聚带；2）采用回剥法并利用最新资料进行校正，得到了研究区更为可靠的构造沉降曲线，重新划分了裂陷期和裂后期的分界，认为32Ma南海海底扩张之后裂陷作用仍在持续，直到23Ma左右才开始大规模裂后热沉降，并进一步解释了裂陷期延迟的形成机制；3）应用非连续拉张模型计算拉张系数的方程计算了研究区的壳幔拉张系数，指出了深水区地幔相对于地壳的优势伸展作用；首次运用平衡剖面技术重建了研究区的构造发育史，计算了各构造期的拉张率和沉积速率，指出研究区新生代整体呈现持续拉张，拉张系数在1.1-1.24之间；4）精细刻画了水合物钻采区的地质构造特征，建立了该区天然气水合物成藏的概念模式；建立了一套根据地震叠加速度计算流体势的方法，为水合物成藏规律的研究提供了新的思路。

Morphology and infraciliature of three species of Metaurostylopsis (Ciliophora, Stichotrichia): M. songi n. sp., M. salina n. sp., and M. marina (Kahl 1932) from sediments, saline ponds, and coastal waters.

Two new urostylid ciliates, Metaurostylopsis songi n. sp. and Metaurostylopsis salina n. sp. and Metaurostylopsis marina (Kahl 1932) are investigated using live observation and protargol impregnation. These species were isolated in Korea from intertidal sediments, saline ponds, and coastal waters. Metaurostylopsis songi is in vivo about 120 pm x 25 mu m, has a slenderly ellipsoidal body, colorless cortical granules in rows on ventral and dorsal body sides, about 54 macronuclear nodules, 28-47 adoral membranelles, five frontal, two or three frontoterminal and six or seven transverse cirri, and 9-12 midventral cirral pairs followed posteriorly by 1-3 single cirri. In vivo M. salina is about 60 pin x 25 mu m, has a pyriform body, colorless cortical granules irregularly arranged, about 45 macronuclear nodules, 18-23 adoral membranelles, three frontal, three to five frontoterminal and two to five transverse cirri, and four or five midventral cirral pairs followed posteriorly by five to seven single cirri. Both species have three marginal cirral rows on each body side and 3 long dorsal kineties. The Korean specimens of M. marina match the Chinese population in all main features. Metaurostylopsis songi differs from M. marina by the more slender body, the number of frontal cirri (invariably five vs. four), and the arrangement of cortical granules (in rows on dorsal and ventral cortex vs. only along dorsal kinetics and anterior body margin). Metaurostylopsis salina differs from its congeners by the distinctly smaller size, the pyriform body shape, the scattered cortical granules (vs. in rows), and number of frontal cirri. It differs from M. marina also by the number of midventral cirral pairs (four or five vs. seven to 11).