华南寒武纪早期刺胞动物演化框架

刺胞动物是华南寒武纪早期海洋生态系统的重要组成部分,针对这类化石的研究有助于深入了解寒武纪多幕式爆发事件。本文主要根据寒武纪早期扬子板块陕南宽川铺生物群、湖北岩家河生物群以及澄江生物群产出的刺胞动物,探讨刺胞动物在寒武纪早期的演化框架,取得了以下初步认识。截至目前,所发现的寒武纪早期刺胞动物绝大多数都属于水母超纲,珊瑚纲的化石记录在寒武系幸运阶、第二阶保存较为罕见。其中寒武系幸运阶刺胞动物的化石记录全部属于底栖类型,体型小,且具有多种类型的身体辐射对称形式,绝大多数类型可以确定属于直接发育。第二阶刺胞动物仍以底栖固着类型为主,体形增大,仅见两、四辐射对称形式;而寒武系第三阶刺胞动物仍有底栖固着类型,但游泳的水母开始出现,体型增大明显,代表水母冠群的出现,以及世代交替的复杂生命周期的正式建立。从幸运阶到第三阶刺胞动物体型明显增大,可能与海水氧气含量增加有关。在寒武纪大爆发的背景下,刺胞动物的分异度和丰度在寒武系幸运阶就已经达到顶峰,然后在第二阶、第三阶开始衰减,这种变化可能与两侧对称动物辐射演化有关。华南刺胞动物的化石记录表明,寒武纪大爆发并非纯粹的一幕式爆发,其中充斥着一系列生物群的替代甚至灭绝事件。     

Extinctions, Morphological Gaps, Major Transitions, Stem Groups, and the Origin of Major Clades, with a Focus on Early Animals

Systematic extinctions can leave major morphological gaps between living crown-group clades. Such morphological gaps would be perceived, from a neontological point of view, as major evolutionary transitions. In order to fill these morphological gaps and to map the evolutionary steps toward major evolutionary transitions, we need to integrate extinct stem-group taxa in phylogenetic studies. However, the recognition of stem group has not been widely adopted in the study of early animal fossils, despite that all fossils are stem groups at one level or another. Part of the difficulty is that stem groups may not have all features that collectively diagnose the respective crown group, and they can have unique (autapomorphic) features, making them tantalizingly similar to and frustratingly different from the crown group (e.g., stem-group eukaryotes can be prokaryotic and stem-group animals can be protistan). The need to embrace stem groups and to implement the PhyloCode, in order to achieve phylogenetic clarity and to offer key paleontological insights into the origin and early animal evolution, is illustrated in debates on several controversial Ediacaran and Cambrian fossils.     

Internal Microanatomy and Zoological Affinity of the Early Cambrian Olivooides

The early Cambrian pentamerous microfossil Olivooides/Punctatus in South China, which is characterized by a diagnostic stellate tubular apex, has been well-known for its almost complete development sequence that can be confidently traced from embryos and hatched juveniles, to conical adults. However, its zoological affinity remains highly controversial. Here we describe the internal microanatomic structures of the soft-body inside the peridermal theca of Olivooides multisulcatus Qian, 1977, including interradial pairs of tentacles, adradial and perradial frenula, perradial oral marginal lappets and twins of perradial gastric saccule-like humps as well as a circular velarium with striated coronal muscles. Particularly, one specimen shows bifurcated velarial canals along the bell aperture. Both the components of the soft-body and the external theca (or cyst) are arranged in perfect pentaradial symmetry. These characteristics are more compatible with those of living cubomedusans and co-occurring Cambrian athecate embryonic cubozoans. Concerning the presence of peridermal theca, Olivooides most likely represents an extinct thecate stem-group cubomedusae but devoid of both perradial eyes and specialized pedalia. The well-grown soft body inside the peridermal tube displays a set of mixed features of both polyp and medusa.     

三峡地区寒武系纽芬兰统岩家河组小壳化石研究进展

埃迪卡拉纪—寒武纪转换时期小壳化石的突然出现是世人瞩目的重要演化事件之一,它见证了动物的爆发式快速演化过程,标志着以两侧对称动物为特征的寒武纪生态系统的开始。湖北三峡地区寒武系纽芬兰统岩家河组处于幸运阶—第二阶这一关键层段,是研究上述问题的理想地区之一。岩家河组小壳化石研究最近取得的进展,为寒武系纽芬兰统幸运阶/第二阶界线的标定,和寒武纪早期软体动物和刺胞动物的演化及个体发育提供了重要的化石依据,本文对其进行了总结。全球界线层型剖面和点位(GSSP)寒武系第二阶底界划分标志的Watsonella crosbyi和Aldanella attleborensis在岩家河组第5段底部的同时出现,意味着岩家河组第5段已归属寒武系第二阶。这两类软体动物化石均广泛分布于华南、蒙古、西伯利亚和北美等大陆碳酸盐岩相寒武系第二阶地层中,它们均是定义寒武系第二阶底界的有力竞争者。另外,W.crosbyi壳顶下方一对肌肉附着结构的发现提供了重要的软组织结构信息,证实了W.crosbyi为内腹型壳,归属为未扭曲的软体动物太阳女神螺类。新发现六方锥石类Septuconularia yanjiaheensis具侧向压扁、两辐射对称的、14个等大锥面的围鞘,是迄今所描述的六方锥石类中锥面数最多的类型。根据横肋疏密程度,围鞘由下向上可分为三部分,可能代表胚胎期、幼年期和成年期不同的生长阶段。Septuconularia显然由寒武系幸运阶Hexaconularia通过扇面扩增演变而来,证明Hexaconularia可能是Arthrochites和Septuconularia之间的中间过渡类型。新发现的橄榄蛋类Octapyrgites elongatus与幸运阶Olivooides和Quadrapyrgites十分相似,围鞘由方形的壳顶区和波纹状倒宝塔形的远顶区组成。然而Octapyrgites只有8个口折叶,有别于具12个口折叶的Quadrapyrgites。通过与幸运期水母类多种对称方式对比,发现在寒武系第二阶橄榄蛋类数量减少和五辐射对称骨状壳类缺失,表明在幸运期/第二期转换时期,底栖固着型的刺胞动物水母类多样性发生了明显衰落,甚至部分绝灭,而同期的两侧对称动物则迅速辐射演化。     

Divergent evolution of medusozoan symmetric patterns: Evidence from the microanatomy of Cambrian tetramerous cubozoans from South China

Living medusozoans and their Middle Cambrian ancestors are characterized fundamentally by four-fold symmetry. The symmetric pattern of their earlier antecedents during the Ediacaran–Cambrian transition, traditionally expected to be tetramerous, needs to be reconsidered in the light of Cambrian pentamerous fossil medusozoans. Here we present a microanatomic analysis of three tiny tetramerous specimens from the Lower Cambrian Kuanchuanpu Formation in southern China; they display diagnostic characteristics of Cubomedusae, including paired but well-separated interradial tentacles deep in the subumbrellar cavity, interradial septa connected by claustra, coronal muscles at the subumbrella, and an annular velarium suspended by twelve frenula. These fossils likely represent three new taxa of a monophyletic stem group of cubomedusans, which is further confirmed by our phylogenetic analysis based on the data matrix with 25 taxa and 107 characteristics. Among these fossil cubozoans and co-occurring pentamerous forms, the specimen ELISN31-31 is mostly close to the crown-group Cubomedusae with regard to its narrow vascular cavity and the endodermic perradial fusion. Diverse symmetrical patterns among different lineages of early Cambrian medusozoan candidates might have been independently evolved in different classes of medusozoans.     

Cambrian explosion: Birth of tree of animals

Excluding the sponges the Kingdom Animalia is usually divided into three subkingdoms: Diploblasta, Protostomia and Deuterostomia. The Cambrian Explosion consists of three major episodes, two of which were in the early Early Cambrian (one represented by the small skeletal fossils “SSFs” at the base of the Cambrian and the other represented by the succeeding Chengjiang faunas “CFs”), and the other episode as their prelude took place in the “Eocambrian” (i.e. the latest Precambrian), represented by the Ediacaran faunas. This unique Big Bang of life has been recognized as giving birth to the entire morphological Tree Of Animals (or metazoans), in short the TOA. Its “seed” in the deep Precambrian, represented by some sort of protist from which the complete TOA must have grown, remains unknown paleontologically. However, the fossil evidence suggests that the three major episodes of the Cambrian Explosion are responsible for the earliest radiations of the three subkingdoms of animals respectively. While the observed Ediacaran fauna might constitute only a small part of the whole Ediacaran biota, our evidence supports that it was dominated by diploblasts (the “trunk” of the TOA) with only a few possible stem-group triploblasts. The Early Cambrian in turn in two phases explosively yielded almost all the major triploblastic crown-branches (Bilateria: the huge “crown” of the TOA), which include the other two subkingdoms: first the extremely diverse protostomes in the Meishucunian Age and then followed by a nearly entire lineage of early deuterostomes from the Chengjiang, including even its most derived member – the earliest true vertebrates. Among the four most significant milestones of morphological origins and radiations in animal history, the first one (i.e. appearance of metazoans) took place in the Ediacaran Period or earlier times, and the other three can be seen in the windows available from the Chengjiang and the Meishucunian fossil assemblages. The newly discovered extinct Phylum Vetulicolia, which has primitively segmented body with simple gill slits in its anterior division, most probably represents one of the roots of the deuterostome subkingdom. Showing a mosaic of basic features possessed in both the bilateral vetulicolians and some primitive echinoderms, the soft-bodied vetulocystids are best regarded as one of the roots of the extant pentamerous echinoderms. Standing on the “top” of the deuterostome super-branch in the early Cambrian TOA are the “the first fish” Myllokunmingia and Haikouichthys, which bear paired eyes and salient proto-vertebrae. These animals represent the real root of the remainder of the vertebrates or craniates. On the contrary, yunnanozoans, including Yunnanozoon and Haikouella, possess neither eyes nor unequivocal vertebrae, and may have nothing to do with the craniates, let alone the vertebrates. Those enigmatic creatures share a similar body-plan with vetulicolians and should be treated as a side-branch within the lower deuterostomes.     

我国古老微体双壳类的发现及其意义

关于双壳类的地质时代延限,前人认为是从奥陶纪或中寒武世至现代。Barrande 和Walcott 将纽约州早寒武世的Fordilla troyensis和Modioloides prisca等属种作为双壳类描述,但Ulrich和Bassler认为该属种是节肢动物门的甲壳类,而非软体动物门的瓣鳃类(双壳类)。     

Morphology and ontogeny of Hunanocephalus ovalis (trilobite) from the Cambrian of South China

The juvenile morphology and ontogeny of the Cheiruroidid trilobite Hunanocephalus ovalis Lee, 1963 from the lower Cambrian Shuijingtuo Formation in Hubei Province, South China is presented. The new material comprises a relatively complete meraspid ontogenetic series (degree 0 to 10), which reveals more details on their morphological changes such as the contraction and disappearance of the pronounced posteromedial notch in the pygidium and the addition of the trunk segments, which are all documented for the first time and can also be used as developmental markers defining their ontogenetic phases. The trunk segmentation schedule of H. ovalis is also discussed, which is similar to the other early Cambrian oryctocephalid trilobites, i.e. as the boundary between the thorax and pygidium migrated posteriorly there is no change in the trunk segment number; the processes of liberation of the thoracic segment and segment insertion into the pygidium are separated from one another, implying that the control of trunk exoskeletal segment appearance and articulation might be decoupled in these trilobites.     

Origin, diversification, and relationships of Cambrian lobopods

As an extinct group, the origin, diversification and relationships of Cambrian lobopods have long been one of the most hotly-debated subjects. Lobopods used to be regarded as closely related to modern onychophorans and tardigrades, but the similarities are based on the few species of Cambrian lobopods and often on a fairly general level. With the discovery of new creatures and based on new observations on fossil lobopods, we consider that Cambrian lobopods show great diversity and reveal a close relationship between Cambrian lobopods and arthropods. In addition, comparison between the dorsal spines of Cambrian lobopods and small shelly fossils suggest that Cambrian lobopods might have originated from the Meishucun radiation and diversified in the Qiongzhusi radiation.     

Early Cambrian palaeogeography and the probable Iberia–Siberia connection

The end of the Proterozoic–beginning of the Cambrian is marked by some of the most dramatic events in the history of Earth. The fall of the Ediacaran biota, followed by the Cambrian Explosion of skeletonised bilaterians, a pronounced shift in oceanic and atmospheric chemistry and rapid climatic change from ‘snowball earth’ to ‘greenhouse’ conditions all happened within a rather geologically short period of time. These events took place against a background of the rearrangement of the prevailing supercontinent; some authors view this as a sequence of individual supercontinents such as Mesoproterozoic Midgardia, Neoproterozoic Rodinia and Early Cambrian Pannotia. Assembled in the Mesoproterozoic, this supercontinent appears to have existed through the Neoproterozoic into the Early Cambrian with periodic changes in configuration. The final rearrangement took place during the Precambrian–Cambrian transition with the Cadomian and related phases of the Pan-African orogeny. The distribution of Early Cambrian molluscs and other small shelly fossils (SSF) across all continents indicates a close geographic proximity of all major cratonic basins that is consistent with the continued existence of the supercontinent at that time. Subsequently, Rodinia experienced breakup that led to the amalgamation of Gondwana, separation of Laurentia, Baltica, Siberia and some small terranes and the emergence of oceanic basins between them. Spreading oceanic basins caused a gradual geographic isolation of the faunal assemblages that were united during the Vendian–Early Cambrian.     

中扬子区寒武纪层序地层研究

笔者建立了中扬子区寒武系层序地层格架，研究了中扬子区层序界面和体系域特点，分析了中扬子区层序样式的基本类型及其特征。共划分出2个二级层序、14个三级层序。寒武系下统划分了1个二级层序(SS3)、6个三级层序(SQ1—SQ6)，平均延时约5．0Ma；体系域颜色普遍较深，泥质含量较高，是一套良好的烃源岩。寒武系中上统划分出1个二级层序(SS4)、8个三级层序(SQ7—SQ14)，平均延时约2．9Ma；体系域颜色普遍较浅，云质含量较高，是一套良好的储集层。总体上H型样式为主，T型样式和TH型样式数量相当，反映寒武纪水体中等。二级层序体系域和三级层序体系域都表现出明显的南北向差异性。     

Lithofacies palaeogeography of the Cambrian in Tarim area

The Tarim area in this paper refers to the present Tarim Basin and its surrounding mountains, bounded on the north by the Middle Tianshan Mountains, on the south by the Kunlun Mountains, on the west by the national boundary of China, and on the east by the Altun Mountains. It covers an area of more than 1 000 000 km². Based on the quantitative and qualitative data from the stratigraphy and petrology of outcrop and well sections, and according to the single factor analysis and multifactor comprehensive mapping method, the single factor maps of the Lower, Middle and Upper Cambrian, and the lithofacies palaeogeography maps of the Early, Middle and Late Cambrian of the Tarim area, are compiled. The most important character of these lithofacies palaeogeography maps is quantification, i.e. the determination of each palaeogeography unit based on accurate quantitative data and the quantitative single factor maps. The study marks the first time that this quantitative method has been applied in the Tarim area. In the Early Cambrian, the present Tarim Basin was mainly a carbonate platform, in which there were penebank, dolostone flats, and gypsum-halite lake. The South Tianshan Basin and East Tarim Basin were siliceous rock-limestone-mudstone basins. The Kunlun Platform was situated in the south, where there were two small lands. The Altun Land was situated in the southeast. In the Middle and Late Cambrian, the framework of lithofacies palaeogeography was similar to that in the Early Cambrian; the land area was gradually reduced; the basin area was enlarged; while the Tarim Carbonate Platform changed little. The transgression continued from the Early Cambrian to the Late Cambrian. Translated from Journal of Palaeogeography, 2006, 8(4): 427–439 [译自: 古地理学报]     

寒武纪三叶虫生物地理区的一些问题

文内叙述寒武纪三叶虫分类、分布和分区。世界上有两个寒武纪生物地理区,一个是北区,另一个是南区。斜坡带的球接子类及少数多节类是世界性分布的,仅能视作任何一区内的生态小区。两区的分界在北非的北部、土耳其的南部,再向东连接天山的北部边缘及蒙古的南部边缘。从南欧寒武纪三叶虫的情况看,是海水深度频繁变化的结果。中国任何一个地区,既不属东冈瓦纳,也不属西冈瓦纳。     

Cambrian stratigraphy of eastern Yunnan

Based on investigation of several types of Cambrian sections in eastern Yunnan, we have tentatively concluded: 1) existence of the Middle Cambrian in eastern Yunnan is definitely proved; 2) the Lower Cambrian of eastern Yunnan may be divided into four formations, of which the Mei-shu-ts'un is so named for the phosphate bearing beds; 3) judging from the lithological character we recognize that the subsiding area of eastern Yunnan is still considered in the late Early Cambrian era; 4) Paragraulos and Redlichia chinensis should not be used as zoning fossils for the Tsanglangpu and Lung-wang-miao formations, respectively; 5) the Middle Cambrian of eastern Yunnan might be correlated with the Middle Cambrian of Kueichow, because Kunming was a subsiding area in the Middle Cambrian era. – Authors.     

塔里木盆地寒武系层序地层格架

通过露头、钻测井及地震层序地层特征综合分析,建立了塔里木盆地寒武系三级层序地层格架,将寒武系划分出6个可全盆地追踪与对比的三级层序。结合反映全球海平面相对变化的全岩碳同位素旋回与反映沉积古水深相对变化的沉积旋回的对比分析,可将塔里木盆地寒武系三级层序分为主要受全球海平面变化控制形成的全球海平面层序(层序CSQ2、CSQ4、CSQ5、CSQ6)及主要受地区性构造沉降作用控制形成的构造层序(层序CSQ1、CSQ3)2类。塔北轮南地区寒武系台地边缘总体呈现由早寒武世的缓坡向中—晚寒武世的镶边台地的演变以及由西向东的进积叠置型式,推测主要与寒武纪全球海平面总体微弱下降及塔里木盆地基底沉降速率总体稳定且较小有关。     

华北克拉通北缘寒武系层序地层划分

华北克拉通北缘寒武系碳酸盐岩作为区域油气勘探的重要目标,其沉积环境演化和层序地层格架的研究对区域地层及有利储层定位具有重要指导意义,也为华北地区寒武系油气勘探提供线索。以大量露头剖面实测分析为基础,结合典型井录井、测井、微观显微镜下分析以及地化测试等数据资料,在年代地层框架下进行旋回性地层分析研究,对华北克拉通北缘寒武系岩性、岩相以及地层叠置样式进行详细剖析,研究沉积趋势变化,总结寒武系层序地层划分样式。研究认为：在全球海平面变化控制下,华北地区自府君山组开始,发育广泛的由南向北的持续海侵,在陆源碎屑供给以及碳酸盐岩加积作用的相互制约下,华北地区表现为碳酸盐台地的形成-发展-镶边过程;以张夏组为界,中-下寒武统表现为缓坡-陆棚的沉积环境,上寒武统为具镶边的台地沉积背景;在不同层位不同层序边界类型（I型、Ⅱ型、Ⅲ型）的分隔作用下,将华北克拉通北缘寒武系划分为7个可区域性对比的三级层序,各三级层序高水位体系域多表现厚层清水碳酸盐岩沉积的特点,且受不同程度白云石化作用,成为区域寒武系重要的有利油气储集层。     

塔里木地区寒武纪岩相古地理

塔里木地区是以当今的塔里木盆地为主体并包括其周边山地的广大地区,北至中天山,南至西昆仑山,西至我国国界,东至阿尔金山,面积逾100万km2。在39个露头剖面和钻井剖面的各种单因素定量及定性资料的基础上,采用单因素分析多因素综合作图法,编制出了塔里木地区寒武系下、中、上统的各种单因素图及相应的寒武纪早、中、晚世的岩相古地理图。这些岩相古地理图最主要的特征是定量,即每个古地理单位的划分和确定都有确切的定量资料与定量的单因素图。这种定量的岩相古地理图在塔里木地区还是首次出现。在早寒武世,塔里木地区是一个碳酸盐岩台地,其中散布着准滩、云坪和膏盐湖。其北侧和东北侧是南天山和塔东硅岩石灰岩泥岩盆地。其南侧是昆仑台地,其中有2个小陆地。其东南侧有阿尔金陆。中寒武世和晚寒武世的岩相古地理格局与早寒武世相似,但陆地面积逐渐缩小,盆地面积逐渐扩大,塔里木碳酸盐岩台地变化不大。从早寒武世到晚寒武世是一个逐渐的海进过程。     

八臂仙母虫(Eoandromeda octobrachiata)新材料及其意义

在中国贵州江口地区陡山沱组新发现的化石材料表明,伊迪卡拉/震旦纪(Ediacaran/Sinian)的宏体化石——八臂仙母虫Eoandromeda octobrachiata的原始生活状态可能为一个较高的圆锥形生命体,具有沿着侧面锥状螺旋排列的八条旋臂.旋臂表面保存有与栉水母动物的栉板可能同源的横向有机质条带,规则排列...     

Mo isotopic variations of a Cambrian sedimentary profile in the Huangling area,South China: Evidence for redox environment corresponding to the Cambrian Explosion

The radiation of macroscopic animals in the Early Cambrian, commonly be called as ‘Cambrian explosion’, had a close relationship with Earth oxygenation. These macrofossils are widely preserved within the Early Cambrian in the northern and western Yangtze Craton. In order to show the casual relationship between animal evolution and Earth redox environment, Mo content and isotopic composition analyses have been conducted on dolomites of the Tianzhushan Formation and the black shales of the Shuijingtuo Formation from the lowermost Cambrian profile in the eastern Huangling dome, northern Yangtze Craton. The calcitic dolostones from the Tianzhushan Formation display lower concentrations of Mo contents (0.055–0.666 ppm) than that of the black silty shales from the Shuijingtuo Formation (6.0–172 ppm). This magnitude difference is most likely related to the different lithologies due to Mo enrichment in shales. The Mo isotope values (δ^98/95Mo) of the calcitic dolostones for the Tianzhushan Formation vary from −0.73‰ to 0.41‰. In comparison, the black sandy shales from the Shuijingtuo Formation exhibit δ^98/95Mo of −1.04–1.84‰, including 5 samples from the earliest Shuijingtuo Formation displaying systematically higher δ^98/95Mo values (0.09–1.84‰) than those of the other 5 samples from upper layers (δ^98/95Mo = −1.04 to −0.19‰). These geochemical data suggest that the redox environment of the early Cambrian is fluctuating, and the peak oxygenation of the ocean had already reached a level similar to modern environments. According to the Mo concentrations and δ^98/95Mo data, the sedimentary succession can be divided into five intervals, which are closely correlated to the episodic evolution of early life in the Cambrian. Together with the available published data from other profiles in the Yangtze Craton, our new data reveal that the higher oxidation environment had a positive effect on the evolution of early life.     

Origin of the Lower Cambrian phosphatic bed at Vassbo, Sweden

An extensive phosphatic bed characterized by a black apatite-cemented sandstone is developed in the uppermost part of the Lower Cambrian formation over the entire Vassbo area, Sweden. The thickness of the layer is generally 0.1 m, increasing in places to 0.2 m. The lower boundary of the bed is transitional to a sandstone; the upper boundary displays a contact that is both sharp and irregular with a Middle Cambrian bituminous black shale.
The phosphatic bed was diagenetically formed by inorganic in situ precipitation of apatite in a sandstone at a depth of about 0.2 m below the sediment-water interface, where reducing conditions were developed, as a result of oxygen consumption during the degradation of organic matter. Sulphur isotope data from associated pyrite suggest that organic processes, such as bacterial sulphate reduction, were also involved in the formation of the phosphatic bed. The depositional environment of the bed was a marine shelf with a zero to very low sedimentation rate. The upper part of the apatite-cemented sandstone was partly reworked by bottom currents as demonstrated by a board containing numerous pebbles. Sedimentological as well as chemical data confirm a southern hemisphere position of the Baltic Shield in the early Cambrian in which an oceanic upwelling and high organic productivity was the probable source of the phosphorus along the palaeo-west-coast.