Lower Cambrian small shelly faunas from Zhejiang (China)and their biostratigraphical implications

Despite a long history of research on the Early Cambrian in China most available data on small skeletal fossils concern fossil associations of the shallow carbonate platform. Information on skeletal fossils from marginal shelf environments of the Yangtze Platform is scanty, which may reflect the rarity of fossils in deeper sedimentary environments but is also due to limitation of carbonate distribution and outcrops, difficulties in fossil extraction, and a general research focus on the Precambrian-Cambrian boundary beds on the carbonate platform. Here we present a documentation of Meishucunian to Qiongzhusian small skeletal fossils from the lower Hetang Formation and the chert unit at its base from the Jiangshan region, Zhejiang Province, representing a relatively deep shelf environment compared to the inner shelf region. The earliest association (Meishucunian) from the chert unit underlying the Hetang Formation is mainly characterized by the occurrence of Protohertzina anabarica, P. unguliformis, Fengzuella zhejiangensis, and Kaiyangites novilis, which differs somewhat in composition from SSF-associations of typical inner shelf deposits. The enigmatic skeletal fossil Fengzuella zhejiangensis, which exhibits an unusual secretional growth mode previously unrecognized from the Early Cambrian, is described in detail. A younger (Qiongzhusian) fossil association contains numerous arthropod remains, such as disarticulated spines of arthropods (Jiangshanodus- and Kijacus-type), which have previously been considered as conodont-like fossils, and bradoriid valves.     

Phylogeny and life habits of Early Arthropods-Predation in the Early Cambrian Sea

We investigated two new arthropods from the Maotianshan-Shale fauna of southern China in the course of our research on life strategies, particularly predation, in Early Cambrian marine macrofaunal biota. One form clearly belongs to the so-called "great-appendage" arthropods, animals that were, most likely, active predators catching prey with their first pair of large, specialized frontoventral appendages. Based on this, we hypothesize that the new species and many others, if not all, of the "great-appendage" arthropods were derivatives of the chelicerate stem lineage and not forms having branched off at different nodes along the evolutionary lineage of the Arthropoda. Rather, we consider the "great-appendage" arthropods as belonging to a monophyletic clade, which modified autapomorphically their first pair of appendages (antennae in general arthropod terminology) into raptorial organs for food capture. The second new form resembles another Maotianshan-Shale arthropod, Fuxianhuia protensa, in sharing a head made of only two separate segments, a small segment bearing oval eyes laterally, and another bearing a large tergite, which forms a wide shield freely overhanging the subsequent narrow trunk segments. This segment bears a single pair of rather short anteriorly directed uniramous appendages, considered as the "still" limb-shaped antennae. Particularly the evolutionary status of head and limbs of these two forms suggests that both are representatives of the early part of the stem lineage toward the crown-group of Arthropoda, the Euarthropoda. These forms appear rather unspecialized, but may have been but simple predators. This adds to our hypothesis that predation was a common, if not dominant feeding strategy in the Cambrian, at least for arthropods.     

Stratigraphic implications of Sinian-Early Cambrian volcanic ash beds on the Yangtze Platform

Volcanic ash beds from shallow- to deep-water facies strata of the Sinian-Early Cambrian (Meishucunian) on the Yangtze Platform consist of bentonites and tuffites which are readily recognized in the field by their physical features and confirmed by geochemical analyses.Geochemistry suggests that the volcanic ash beds in Meishucunian time are rhyolite and rhyodacite while those in the Qiongzhusian and Sinian are andesite and trachyandesite.The ash beds in the time-equivalent strata, even in different areas display rather similar geochemical features, whereas the ash beds in different strata even in the same areas show large chemical difference.The results suggest that these ash beds can be used for intra- and extra-basinal correlations of the Sinian-Early Cambrian interval on the Yangtze Platform.Additionally, these ash beds suggest high potentials for further U-Pb dating strategies.     

Phosphatized embryo fossils from the Cambrian explosion

The earliest Cambrian (Meishucunian) phosphatized metazoanOlivooides from the Yangtze platform is represented by dierent developmental stages: from blastula through possible gastrula to tissue differentiation and finally to the hatched animals. The fossilization of embryos opens a new field in palaeontologypalaeoembryology. Embryological studies will be of importance for the understanding of ontogeny and phylogeny of metazoans from the Cambrian explosion.     

Yangtzedonta and the early evolution of shelled molluscs

The holotype of Yangtzedonta Yu, 1985 and its type species, Y. primitiva Yu, 1985, is an incomplete specimen of Xianfengella prima He et Yang, 1982. Therefore, based on the principle of priority, the names Yangtzedonta and Y. primitiva Yu (1985) should be abandoned. The fact that ventral valve of Xianfengella is bilaterally symmetrical indicates that they are not the left valve of a bivalved mollusc, as originally proposed by Yu. At present, the oldest certain bivalve is Xianfengoconcha elliptica Zhang, which occurs in the Early Cambrian Qiongzhusi Stage in China. The oldest known rostroconch is Heraultipegma yunnanensis He et Yang, which first occurs in the third small shelly fossil assemblage zone of the Early Cambrian Meishucunian Stage. The oldest known monoplacophoran, Maikhanella pristinis (Jiang), first occurs in the first small shelly fossil assemblage zone (Meishucunian Stage). Evidence derived from analyses of shell morphology, mode of growth, histology, and stratigraphic occurrence suggests that bivalves evolved from a branch of rostroconchs, and that rostroconchs in turn evolved from a branch of monoplacophorans. The hypothesis that these three groups originated simultaneously and independently at the beginning of the Cambrian and subsequently evolved in parallel, probably is incorrect.     

Possible developmental mechanisms underlying the origin of the crown lineages of arthropods

The extraordinarily preserved, diverse arthropod fauna from the Lower Cambrian Maotianshan shale, central Yunnan (southwest China), represents different evolutionary stages stepping from stem lineages towards crown arthropods (also called euarthropods), which makes this fauna extremely significant for discussion of the origin and early diversification of the arthropods. Anatomical analyses of the Maotianshan shale arthropods strongly indicate that     

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.     

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.     

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.     

Sophisticated particle-feeding in a large Early Cambrian crustacean

Most Cambrian arthropods employed simple feeding mechanisms requiring only low degrees of appendage differentiation. In contrast, post-Cambrian crustaceans exhibit a wide diversity of feeding specializations and possess a vast ecological repertoire. Crustaceans are evident in the Cambrian fossil record, but have hitherto been known exclusively from small individuals with limited appendage differentiation. Here we describe a sophisticated feeding apparatus from an Early Cambrian arthropod that had a body length of several centimetres. Details of the mouthparts resolve this taxon as a probable crown-group (pan)crustacean, while its feeding style, which allowed it to generate and handle fine food particles, significantly expands the known ecological capabilities of Cambrian arthropods. This Early Cambrian record predates the major expansions of large-bodied, particle-handling crustaceans by at least one hundred million years, emphasizing the importance of ecological context in driving adaptive radiations.     

A Cambrian micro-lobopodian and the evolution of arthropod locomotion and reproduction

The evolutionary success of arthropods, the most abundant and diverse animal group, is mainly based on their segmented body and jointed appendages, features that had evolved most likely already before the Cambrian. The first arthropod-like animals, the lobopodians from the Early Cambrian, were unsclerotized and worm-like, and they had unjointed tubular legs. Here we describe the first three-dimensionally preserved Cambrian lobopodian. The material presented of Orstenotubulus evamuellerae gen. et sp. nov. is the smallest and youngest of a lobopodian known. O. evamuellerae shows strikingly detailed similarities to Recent tardigrades and/or onychophorans in its cellular-structured cuticle and the telescopic spines. It also shows similarities to other, longer known lobopodians, but which are ten times as large as the new form. These similarities include the finely annulated body and legs, which is characteristic also for Recent onychophorans, and paired humps continuing into spines situated dorsally to the leg insertions, a feature lacking in the extant forms. The morphology of O. evamuellerae not only elucidates our knowledge about lobopodians, but also aids in a clearer picture of the early evolution of arthropods. An example is the single ventral gonopore between a limb pair of O. evamuellerae, which indicates that a single gonopore, as developed in onychophorans, tardigrades, pentastomids, myriapods and insects, might represent the plesiomorphic state for Arthropoda, while the paired state in chelicerates and crustaceans was convergently achieved. Concerning life habits, the lateral orientation of the limbs and their anchoring spines of the new lobopodian imply that early arthropods were crawlers rather than walkers.     

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.     

分布式多目标跟踪中数据融合的一种新方法

数据融合问题是分布式多传感器多目标跟踪中的关键问题。相关文献在航迹和假设的相容性判别以及数据融合过程的分析等问题上存在一些不足。该文从“无矛盾原则”出发,给出了准确而完善的航迹和假设相容性判别条件,澄清了前人在此问题上的一些模糊概念。对航迹和假设的融合过程做了详细的分析,提出了“定向融合航迹”和“自由融合航迹”的概念。按照新的航迹和假设相容性判别条件,进行了仿真实验。实验结果证实新方法是有效性的。     

Acute vision in the giant Cambrian predator Anomalocaris and the origin of compound eyes

Until recently, intricate details of the optical design of non-biomineralized arthropod eyes remained elusive in Cambrian Burgess-Shale-type deposits, despite exceptional preservation of soft-part anatomy in such Konservat-Lagerst?tten. The structure and development of ommatidia in arthropod compound eyes support a single origin some time before the latest common ancestor of crown-group arthropods, but the appearance of compound eyes in the arthropod stem group has been poorly constrained in the absence of adequate fossils. Here we report 2-3-cm paired eyes from the early Cambrian (approximately 515 million years old) Emu Bay Shale of South Australia, assigned to the Cambrian apex predator Anomalocaris. Their preserved visual surfaces are composed of at least 16,000 hexagonally packed ommatidial lenses (in a single eye), rivalling the most acute compound eyes in modern arthropods. The specimens show two distinct taphonomic modes, preserved as iron oxide (after pyrite) and calcium phosphate, demonstrating that disparate styles of early diagenetic mineralization can replicate the same type of extracellular tissue (that is, cuticle) within a single Burgess-Shale-type deposit. These fossils also provide compelling evidence for the arthropod affinities of anomalocaridids, push the origin of compound eyes deeper down the arthropod stem lineage, and indicate that the compound eye evolved before such features as a hardened exoskeleton. The inferred acuity of the anomalocaridid eye is consistent with other evidence that these animals were highly mobile visual predators in the water column. The existence of large, macrophagous nektonic predators possessing sharp vision--such as Anomalocaris--within the early Cambrian ecosystem probably helped to accelerate the escalatory 'arms race' that began over half a billion years ago.     

A Chengjiang-type fossil assemblage from the Hongjingshao Formation （Cambrian Stage 3） at Chenggong,Kunming, Yunnan

A new Chengjiang-type fossil assemblage is reported herein from the lower part of the Hongjingshao Formation at Xiazhuang village of Chenggong,Kunming,Yunnan.The fossil assemblage,named as Xiazhuang fossil assemblage,yields predominantly soft-bodied fossils,including arthropods,brachiopods,priapulids,lobopods and some problematic taxa,with arthropods being the most dominant group.Preservation and composition of the fossil assemblage are very similar to the typical Chengjiang biota,which is preserved in the middle Yu’anshan Formation in the large area of eastern Yunnan.The associated trilobites demonstrate that the soft-bodied fossil assemblage belongs to the late Qiongzhusian in age(Stage 3,Cambrian),suggesting that the Hongjingshao Formation is probably a diachronous lithostratigraphic unit ranging from the upper Qiongzhusian to the lower Canglangpuan stages in eastern Yunnan.The fossil assemblage from the Xiazhuang area fills up the missing link between the typical older Chengjiang biota and the younger Malong and Guanshan biotas,making eastern Yunnan a unique area in the world to reveal the early evolutionary history of animals and palaeocommunity dynamics during the‘‘Cambrian explosion’’.     

Comparative study of Cambrian lobopods <Emphasis Type="Italic">Miraluolishania</Emphasis> and <Emphasis Type="Italic">Luolishania</Emphasis>

The rare fossil Miraluolishania described by Liu et ah from the Lower Cambrian Chengjiang Lagerstatte in 2004 is regarded as an arthropod sphinx because it bears mosaic features of both Iobopods and arthropods. The discovery of this rare transitional form offers direct fossil evidence for exploring the relationship between Iobopods and arthropods. However, some scientists consider Miraluolishania to be a junior synonym of Luolishania because the former superficially resembles the latter in general appearance. Considering the significant differences between the two taxa, a thorough comparative study of Miraluolishania and Luolishania leads to the conclusion that there are definitely two different genera. Nevertheless, the ＂Luolishania＂ of the Haikou area is indeed ＂Miraluolishania＂, whereas Luolishania is most likely the typical genus of the Maotianshan area of Chengjiang County.     

An armoured Cambrian lobopodian from China with arthropod-like appendages

Cambrian fossil Lagerst?tten preserving soft-bodied organisms have contributed much towards our understanding of metazoan origins. Lobopodians are a particularly interesting group that diversified and flourished in the Cambrian seas. Resembling 'worms with legs', they have long attracted much attention in that they may have given rise to both Onychophora (velvet worms) and Tardigrada (water bears), as well as to arthropods in general. Here we describe Diania cactiformis gen. et sp. nov. as an 'armoured' lobopodian from the Chengjiang fossil Lagerst?tte (Cambrian Stage 3), Yunnan, southwestern China. Although sharing features with other typical lobopodians, it is remarkable for possessing robust and probably sclerotized appendages, with what appear to be articulated elements. In terms of limb morphology it is therefore closer to the arthropod condition, to our knowledge, than any lobopodian recorded until now. Phylogenetic analysis recovers it in a derived position, close to Arthropoda; thus, it seems to belong to a grade of organization close to the point of becoming a true arthropod. Further, D. cactiformis could imply that arthropodization (sclerotization of the limbs) preceded arthrodization (sclerotization of the body). Comparing our fossils with other lobopodian appendage morphologies--see Kerygmachela, Jianshanopodia and Megadictyon--reinforces the hypothesis that the group as a whole is paraphyletic, with different taxa expressing different grades of arthropodization.     

The occurrence of the genus Marrella(Trilobitoidea) in Asia

An important discovery in the Kaili Biota has been made recently, and many interesting components from ChengjiangBiota and Burgess Shale Biota have been discovered. Among them Marrella, only known from mid-Middle Cambrian Burgess Shale Biotaof British Columbia, Canada, is one of the most important arthropods fossils. Because the Kaili Biota is older than the Burgess Shale Biota,the strange head shield of Marrella occurring in the Kaili Biota outside Laurentia ranges from mid-Middle Cambrian to early Middle Cam-brian and is significant for the reconstruction of palaeogeography and evolutionary study of early metazoa. In the present paper Marrellasp. is reported for the first time in Asia.