Progress and significance in research on the early Middle Cambrian Kaili Biota, Guizhou Province,China

Important progress in research on the Kaili Biota has been made recently. Many interesting components from Chengjiang Biota and Burgess Shale Biota have been discovered, e.g. Microdictyon of lobopodia; Ottoia, Palaeoscolex of worms; Naraoia, Marrella of Trilobitioidea, Mollisonia, anamalocarids and other non-trilobite arthropods; and new sorts of echinoder-mas, macroalage fossils and so on. Recent work on the Kaili Biota has resulted in the following developments: (i) an increase in the number of animal genera, up to more than 100 genera in total, so that the Kaili Biota has become the third most diverse of the Burgess Shale-type Biota after the Burgess Shale and Chengjiang Biotas; and (ii) the most noteworthy fossils in the Kaili Biota are echinoderms, non-trilobite arthropods and soft-bodied medusiform fossils, especially the most diverse echinoderms. The progress provides envidence for the biodiversity of marine organisms presented after the "Cambrian Explosion" and serves as a link between the earlist Cambrian Chengjiang Biota and late early Middle Cambrian Burgess Shale Biota. It is of great significance in the reconstruction of the Cambrian palaeoplate, palaeongeography and in research on taphonomy.     

凯里动物群与我国早期后生动物演化的初步研究

作者所发现的中寒武世凯里动物群,属于带软躯体化石的布吉斯页岩型的动物群,是继云南澄江动物群之后我国发现的又一个早期后生动物群。它和新近发现的晚震旦世庙河生物群,分别填补了晚震旦世、中寒武世我国带软躯化石生(动)物群的空白,形成了我国多时期带软躯化石的生(动)物群体系。对于研究我国及世界早期后生动物演化并不断揭示其演化规律有重要意义。     

A giant Ordovician anomalocaridid

Anomalocaridids, giant lightly sclerotized invertebrate predators, occur in a number of exceptionally preserved early and middle Cambrian (542-501?million years ago) biotas and have come to symbolize the unfamiliar morphologies displayed by stem organisms in faunas of the Burgess Shale type. They are characterized by a pair of anterior, segmented appendages, a circlet of plates around the mouth, and an elongate segmented trunk lacking true tergites with a pair of flexible lateral lobes per segment. Disarticulated body parts, such as the anterior appendages and oral circlet, had been assigned to a range of taxonomic groups--but the discovery of complete specimens from the middle Cambrian Burgess Shale showed that these disparate elements all belong to a single kind of animal. Phylogenetic analyses support a position of anomalocaridids in the arthropod stem, as a sister group to the euarthropods. The anomalocaridids were the largest animals in Cambrian communities. The youngest unequivocal examples occur in the middle Cambrian Marjum Formation of Utah but an arthropod retaining some anomalocaridid characteristics is present in the Devonian of Germany. Here we report the post-Cambrian occurrence of anomalocaridids, from the Early Ordovician (488-472?million years ago) Fezouata Biota in southeastern Morocco, including specimens larger than any in Cambrian biotas. These giant animals were an important element of some marine communities for about 30?million years longer than previously realized. The Moroccan specimens confirm the presence of a dorsal array of flexible blades attached to a transverse rachis on the trunk segments; these blades probably functioned as gills.     

贵州寒武系棘皮动物研究新进展及其组合特征

贵州寒武系含有丰富的棘皮动物化石,其中凯里组及杷榔组中的始海百合数以千计.近期贵州下司淑里村杷榔组上部粉砂质泥岩中发现的新类型殷氏原始球状始海百合（新属、新种）Protogloboeocrinus yini （gen.et sp.nov.）,使贵州寒武系棘皮动物分异度加大,达8属、种及1科、属、种未定,分属3个亚门.演化趋势是：类群增多,始海百合个体变小,缝孔变圆,吸盘附着的化石类型增多,生态日趋复杂.凯里生物群也是棘皮动物大辐射的窗口.     

Primitive deuterostomes from the Chengjiang Lagerst?tte (Lower Cambrian, China).

Cambrian fossil-Lagerst?tten (sites of exceptional fossil preservation), such as those from Chengjiang (Lower Cambrian) and the Burgess Shale (Middle Cambrian), provide our best window into the Cambrian 'explosion'. Such faunas are known from about 40 localities, and have yielded a widely disparate series of taxa ranging from ctenophores to agnathan fish. Recent excavations of the Chengjiang fossil-Lagerst?tte, known from a series of sites near Kunming in Yunnan, south China, have resulted in the discovery of several new forms. In conjunction with material described earlier, these provide evidence for a new group of metazoans, the vetulicolians. Several features, notably a series of gill slits, suggest that this group can throw light on an early stage of deuterostome diversification.     

贵州黔东地区中-下寒武统凯里组发育特征

贵州黔东地区中－下寒武统凯里组主要由粉砂质泥岩，页岩及灰岩组成，富产三叶虫，其中台江八郎乌溜－曾家岩的凯时组含著名的中寒武世凯里生物群、早寒武世台江生物群，2001年成为国际中－下寒武统层型界线候选部，对其深入研究具有重大科学和实用意义。就我省黔东地区凯里组的岩石地层，生物地层，年代地层，沉积环境等进行研究及总结，亦确认黔东凯里组是一个十分重要的地层单位。     

Proposal and prospects for the global Lower-Middle Cambrian boundary

In recent years paleontologists have been working on the global stratotype section and standard point (GSSP) for the Lower-Middle Cambrian boundary in North America, China, Morocco, Spain and Russia. Continually, they suggest the potential first appearance datum (FAD) of the Middle Cambrian, such as Oryctocephalus indicus, Ovatoryctocara granulata, Acadoparadoxides mureroensis, Hupeolenus and Arthricocephalus chauveaui as being relevant to the potential stratotype section for the Lower-Middle Cambrian boundary. There are only a few trilobites distributed widely enough in the Early to Middle Cambrian connection, so virtually no trilobite(s) can act as the FAD for the global Middle Cambrian. The easily identified Oryctocephalus indicus is widely distributed in three realms of the global Cambrian. Its first appearance position is linked to the extinction of old species coevolution of new ones. It appears to be a comparatively good FAD of Middle Cambrian. The continuous and unbroken Wuliu section (Balang, Taijiang County, Guizhou Province, China) includes O. indicus and co-occurs with many benthic and nektonic trilobites. The evolution of acritarchs in the section, the changed characteristics of trace elements (REE and C isotopes) are generally consistent with the evolution of trilobites both above and below the boundary. Strata above the boundary yield the famous Kaili Biota. The Wuliu section is easily accessible from a main highway.     

贵州台江早寒武世台江动物群的发现及意义

贵州台江八郎凯里组剖面中下部中、下寒武统界线之下厚达１５米左右的粉砂质泥岩、页岩中存在着一个类似凯里动物群的台江动物群,由８个门类３９个属组成,核心为蠕形动物,三叶虫数量很多。形成于内陆棚浅海环境。时代为早寒武世末期,介于著名的早寒武世早期澄江动物群和中寒武世早期凯里动物群之间,对于早期后生动物的演化研究具有重要的意义。     

Discovery of Corallina fossil from the Middle Cambrian of Taijiang County, Guizhou Province, China

Abundant and well-preserved macroalgae fossils have been discovered in silty mudstones and shales from the Middle Cambrian of Taijiang County, Guizhou Province, China. CorallinaParaamphiroa siniansis gen. et sp. nov. in the Kaili Fm., is the earliest Corallina fossil, which usually coexists with other carbonaceous macroalgae fossils on the bedding planes of mudstone and shale and is different from the calcareous and phosphatized Precambrian-Cambrian thallus discovered in the Lower Cambrian of Siberia and the Doushantou phosphate rocks of Weng’an County, Guizhou Province, China. The discovery of calcified thallus in the Kaili Fm. bears great importance on the study of phylogeny and evolution of early calcareous macroalgae.     

A soft-bodied mollusc with radula from the Middle Cambrian Burgess Shale

Odontogriphus omalus was originally described as a problematic non-biomineralized lophophorate organism. Here we re-interpret Odontogriphus based on 189 new specimens including numerous exceptionally well preserved individuals from the Burgess Shale collections of the Royal Ontario Museum. This additional material provides compelling evidence that the feeding apparatus in Odontogriphus is a radula of molluscan architecture comprising two primary bipartite tooth rows attached to a radular membrane and showing replacement by posterior addition. Further characters supporting molluscan affinity include a broad foot bordered by numerous ctenidia located in a mantle groove and a stiffened cuticular dorsum. Odontogriphus has a radula similar to Wiwaxia corrugata but lacks a scleritome. We interpret these animals to be members of an early stem-group mollusc lineage that probably originated in the Neoproterozoic Ediacaran Period, providing support for the retention of a biomat-based grazing community from the late Precambrian Period until at least the Middle Cambrian.     

Eocrinoids from the Cambrian Mantou Formation of Dalian, Liaoning

Cambrian echinoderms,especially eocrinoids,have been reported recently from Guizhou and Yunnan provinces of China,such as the Kaili,Balang,and Guanshan faunas.The present paper reports abundant well-preserved eocrinoids from the Cambrian Mantou Formation of Dalian,Liaoning Province.This new discovery provides additional data on the early evolution of Echinodermata.It also gives us a clue as to where to seek the Cambrian soft-bodied fauna on the margin of the North China Platform.     

下寒武统宽背虫(Bathynotus)的灭绝过程——以贵州剑河八郎凯里组的Bathynotus kueichouensis为例

宽背虫(Bathynotus)这一传统早寒武世或寒武纪第2世晚期最重要的三叶虫属,广泛分布于全球寒武纪3个生物大区,为区域地层对比的标准化石.Bathynotus在贵州剑河八郎乌溜-曾家崖剖面凯里组下部极为丰富.该剖面已成为全球寒武系第2统和第3统界线层型候选剖面,表明凯里组是一个穿时的地层单元,其3个三叶虫带最下部一个带为Bathynotus holopygus-Ovatoryctocara granulata.经过逐层大量的化石标本采集统计证实,乌溜-曾家崖剖面宽背虫(Bathynotus)在距凯里组底部51.60 m处骤然绝灭,距寒武系第2统的顶界(距凯里组底52.80 m)只有1.20 m.本文就乌溜-曾家崖剖面凯里组下部宽背虫(Bathynotus)的数量分布变化认为其灭绝过程是逐步的.     

New data regarding acritarch biostratigraphy from the Early-Middle Cambrian Kaili Formation in Chuandong, Guizhou Province, China

Previous studies have evaluated acritarchs from the Balang and Pingzhai sections of the Early-Middle Cambrian Kaili Formation in east Guizhou, China. The results of these studies suggest that the acritarchs in these sections have potential biostratigraphic significance. More recently, 34 samples collected from 83m lower part of the Lower-Middle cambrian Kaili Formation in the Jianshan section of Chuandong, Guizhou Province were prepared for palynological analysis. Analysis of these samples revealed a distinct change in the acritarch assemblages at the bed, which was approximately 46m above the bottom of the Kaili Formation. These findings suggest a boundary represented by an important alteration of the ecological environment. In addition, the position of the acritarch biostratigraphic change is somewhat higher than the boundary between Cambrian Series 2 and Series 3 based on trilobites found in the same section. This indicates that the distinct change in acritarch assemblages is similar to the change in trilobite assemblages that occurs around the boundary between Cambrian Series 2 and Series 3. Therefore, acritarch biostratigraphy can provide data that can be used to define the base of Cambrian Series 3 in this region, and possibly worldwide.     

Discovery of sponge body fossils from the late Meishucunian (Cambrian) at Jinsha, Guizhou, south China

Here we report discovery of a sponge body fossil Triticispongia sp. from the base of lower Cambrian Niutitang Formation at Jinsha, Guizhou. Stratigraphically, the fossil horizon is located below Ni-Mo ore layer with the Niutitang Biota above, and is equivalent to the late Meishucunian. The species is global in shape with skeletons composed of stauractins and monaxons. Triticispongia sp. reported here may be the earliest sponge body fossils of Cambrian, which provides new information for understanding early evolution and radiation of sponge animals.     

A palaeontological solution to the arthropod head problem

The composition of the arthropod head has been one of the most controversial topics in zoology, with a large number of theories being proposed to account for it over the last century. Although fossils have been recognized as being of potential importance in resolving the issue, a lack of consensus over their systematics has obscured their contribution. Here, I show that a group of previously problematic Cambrian arthropods from the Burgess Shale and Chengjiang faunas form a clade close to crown-group euarthropods, the group containing myriapods, chelicerates, insects and crustaceans. They are characterized by modified or even absent endopods, and two pre-oral appendages. Comparison with reconstructions of the crown-group euarthropod ground plan and recent investigations into onychophorans demonstrates that these two appendages are the first antenna (of extant crustaceans) and a more anterior appendage associated with an ocular segment. The latter appendage has been reduced in all crown-group euarthropods. Its most likely relic is as a component of the labrum. These fossils thus tie together results from disparate living groups (onychophorans and euarthropods).     

华南布尔吉斯页岩型软躯体化石生物群对比研究

目的梳理华南布尔吉斯页岩型软躯体化石生物群的产出层位、生物面貌和沉积环境。方法对10个华南布尔吉斯页岩型软躯体化石生物群进行对比研究。结果除了岩家河生物群之外的9个布尔吉斯页岩型生物群代表了一系列生活在泥砂质基底的、从时空分布上逐渐过渡的具有高度相关性的生物群落。结论布尔吉斯页岩型化石生物群在中晚寒武世应该会有更大规模的分布。     

多枝黄山藻(Huangshanophyton fluticulosum Chen,Lu and Xiao,1994)在凯里生物群中的发现

蓝田植物群处于宏观藻类第一个繁盛期的前期,化石类型丰富,属、种众多,形态结构复杂,具有明显的器官分化,是一个独具特色的植物群。其中多枝黄山藻Huangshanophyton fluticulosum Chen,Lu&Xiao,1994是最常见的分子之一。凯里生物群中的Huangshanophyton fluticulosum,与安徽蓝田组的模式标本相比,藻体形态一致,仅有细微差异,即扇形发散角较大,藻丝直径较小。从埃迪卡拉纪陡山沱期至寒武纪第3世(原中寒武世),经历了近5千多万年特别是埃迪卡拉纪末期的生物灭绝事件后,Huangshanophyton fluticulosum不仅安全渡过而且形态结构特征没有发生多少变化,说明该类生物具有较强的生存竞争能力,以及遗传上的稳定性、演化上的保守性和较强的环境适应性。     

An Early Cambrian tunicate from China

Like the Burgess Shales of Canada, the Chengjiang Lagerst?tte from the Lower Cambrian of China is renowned for the detailed preservation as fossils of delicate, soft-bodied creatures, providing an insight into the Cambrian explosion. The fossils of possible hemichordate chordates and vertebrates have attracted particular attention. Tunicates, or urochordates, comprise the most basal chordate clade, and details of their evolution could be important in understanding the sequence of character acquisition that led to the emergence of chordates and vertebrates. However, definitive fossils of tunicates from the Cambrian are scarce or debatable. Here we report a probable tunicate Cheungkongella ancestralis from the Chengjiang fauna. It resembles the extant ascidian tunicate genus Styela whose morphology could be useful in understanding the origin of the vertebrates.     

Modern optics in exceptionally preserved eyes of Early Cambrian arthropods from Australia

Despite the status of the eye as an "organ of extreme perfection", theory suggests that complex eyes can evolve very rapidly. The fossil record has, until now, been inadequate in providing insight into the early evolution of eyes during the initial radiation of many animal groups known as the Cambrian explosion. This is surprising because Cambrian Burgess-Shale-type deposits are replete with exquisitely preserved animals, especially arthropods, that possess eyes. However, with the exception of biomineralized trilobite eyes, virtually nothing is known about the details of their optical design. Here we report exceptionally preserved fossil eyes from the Early Cambrian (～ 515 million years ago) Emu Bay Shale of South Australia, revealing that some of the earliest arthropods possessed highly advanced compound eyes, each with over 3,000 large ommatidial lenses and a specialized 'bright zone'. These are the oldest non-biomineralized eyes known in such detail, with preservation quality exceeding that found in the Burgess Shale and Chengjiang deposits. Non-biomineralized eyes of similar complexity are otherwise unknown until about 85 million years later. The arrangement and size of the lenses indicate that these eyes belonged to an active predator that was capable of seeing in low light. The eyes are more complex than those known from contemporaneous trilobites and are as advanced as those of many living forms. They provide further evidence that the Cambrian explosion involved rapid innovation in fine-scale anatomy as well as gross morphology, and are consistent with the concept that the development of advanced vision helped to drive this great evolutionary event.