The lobopodian Paucipodia inermis from the Lower Cambrian Chengjiang fauna, Yunnan, China

New specimens of Paucipodia inermis Chen, Zhou & Ramsköld, 1995, are described from the Lower Cambrian Chengjiang Lagerstätte in Haikou, Kunming. Details not previously seen in the Chengjiang material appear to be caused by early diagenetic processes. Some features not previously observed in Palaeozoic lobopodians include details of the dermomuscular sac, body cavities, contents of the gut, possible paired ventral nerve ganglia, and a rasping or biting apparatus with teeth. The latter implies a fundamental difference from onychophorans and rules out an ancestral position for Palaeozoic lobopodians. The supposed tail is shown to be the head, and it is shown that this animal possessed nine pairs of lobopods rather than six, as originally stated. The family Paucipodiidae n. fam. is introduced.     

The earliest myodocopes: ostracodes from the late Ordovician Soom Shale Lagerstätte of South Africa

The late Ordovician Soom Shale Lagerstätte of South Africa has yielded Myodoprimigenia fistuca n. gen. and n. sp., the earliest and only known Ordovician occurrence of myodocopes, one of the major groups of ostracodes. M. fistuca is a likely sister group of the Upper Silurian 'cypridinid' myodocopes and allied forms. It had a thin, lightly mineralized and flexible shell with microstructures resulting from in vivo calcification processes. It probably fed on cephalopod carrion, thus extending evidence for a carnivorous scavenging lifestyle in ostracodes back by 200  Ma. The species was probably nektobenthic and thus consistent with the notion that the origin of the late Silurian pelagic myodocopes - and therefore of pelagic ostracodes - is to be charted in a benthic to pelagic ecological shift in the group.     

A NEW ARTHROPOD IN CHAIN-LIKE ASSOCIATIONS FROM THE CHENGJIANG LAGERSTÄTTE (LOWER CAMBRIAN), YUNNAN, CHINA

Abstract:  A new arthropod, Synophalos xynos gen. et sp. nov., is described from the Lower Cambrian of the Chengjiang Lagerstätte, Yunnan, China. It is provisionally assigned to Crustacea, Waptiida and Waptiidae. Just one example of an isolated individual of S .  xynos is known. Typically, individuals of the new species occur together in similar, robustly integrated chain-like associations. Such a strongly linked configuration is apparently unique for any arthropod, fossil or living. The chain-like association is suggested to represent an example of collective behaviour, possibly associated with reproduction, or more likely migration.     

Moulting in the lobopodian Onychodictyon from the lower Cambrian of Greenland

A number of lobopodian taxa from the Cambrian display pairs of sclerotized plates symmetrically positioned along the dorsum of the animal, predominantly above the walking appendages. Most genera were described from complete body fossils exquisitely preserved in the famous Cambrian Lagerstätten, but lobopodian phosphatized plates are found worldwide as typical components of Cambrian small shelly fossil assemblages (SSF). Details regarding intraspecific and ontogenetic variation in lobopod plates are elusive, and the lack of details of ornamentation in Lagerstätte specimens does not minimize the problem. We document here an assemblage of well‐preserved isolated plates of Onychodictyon sp. from the Lower Cambrian (Cambrian Series 2, Stage 4) of North Greenland. Two specimens exhibit perfectly conjoined plates from successive moults. Details of ornamentation and the outline and profile of the fixed plates are identical, but width and length of the underlying plate are 24% larger. These specimens boost the body of evidence that lobopodians moulted but also show that plate outline and ornamentation did not vary during ontogeny.     

A new shell-bearing organism from the Cambrian Spence Shale of Utah

A new shell-bearing organism with preserved soft tissue, Armilimax pauljamisoni n. gen. n. sp., is reported from the middle Cambrian (Miaolingian: Wuliuan) Miners Hollow locality of the Spence Shale of northern Utah. The described organism is known from a single articulated specimen and preserves a prominent shell, a slug-like body, as well as a U-shaped digestive tract. Its overall appearance is similar to halkieriids, but it does not preserve sclerites. The possible affinities of the new taxon and potential reasons for the presence of a U-shaped gut are discussed. Armilimax pauljamisoni is the first shell-bearing animal of its kind from the Great Basin and extends the diversity of body plans in the Spence Shale Fossil-Lagerstätte.     

A New Bivalved Arthropod from the Early Cambrian Sirius Passet Fauna, North Greenland

A new bivalved arthropod is described from the Lower Cambrian (?Upper Atdabanian) Buen Formation of North Greenland. Pauloterminus spinodorsalis gen. et sp. nov. possesses a bivalved carapace that covers the head, which has a single pair of antennae, and anteriormost thorax. No mouthparts are visible. The five‐segmented abdomen was limbless and terminated in a telson plus a pair of large, lobate uropods. A suite of at least six biramous thoracic limbs are present: the short endopods are made up of small, serial podomeres, while the exopods are lobate and may have functioned as gills as well as in swimming. Partially infilled guts are occasionally visible, suggesting that this animal may have been a sediment feeder. It is compared to other Cambrian bivalved arthropods, especially the waptiids Chuandianella ovata from the Lower Cambrian Chengjiang fauna (China) and Waptia fieldensis from the Middle Cambrian Burgess Shale (British Columbia). Of these three animals, the Greenland and Chinese taxa appear to be the most closely related. P. spinodorsalis possesses many typical arthropod features, but it also demonstrates more primitive characters that are more reminiscent of the lobopodians.     

Anatomy and lifestyles of Early Cambrian priapulid worms exemplified by Corynetis and Anningvermis from the Maotianshan Shale (SW China)

Accurate information on the anatomy and ecology of worms from the Cambrian Lagerstätten of SW China is sparse. The present study of two priapulid worms Anningvermis n. gen. and Corynetis Luo & Hu, 1999 from the Lower Cambrian Maotianshan Shale biota brings new information concerning the anatomical complexity, functional morphology and lifestyles of the Early Cambrian priapulids. Comparisons are made with Recent priapulids from Sweden (live observations, SEM). The cuspidate pharyngeal teeth of Anningvermis (circumoral pentagons) and the most peculiar radiating oral crown of Corynetis added to the very elongate pharynx of these two forms are interpreted as two different types of grasping apparatus possibly involved in the capture of small prey. Corynetis and Anningvermis are two representative examples of the Early Cambrian endobenthic communities largely dominated by priapulid worms (more than ten species in the Maotianshan Shale biota) and to a much lesser extent by brachiopods. Corynetis and Anningvermis were probably active mud-burrowers and predators of small meiobenthic animals. Likewise predator priapulid worms exploited the interface layer between the seawater and bottom sediment, where meiobenthic organisms were abundant and functioned as prey. This implies that complex prey-predator relationship between communities already existed in the Early Cambrian. This study also shows that the circumoral pentagonal teeth and caudal appendage were present in the early stages of the evolutionary history of the group and were important features of the priapulid body plan already in the Early Cambrian. Two new families, one new genus and new species are introduced and described in the appendix.     

MYGALOMORPH SPIDERS (ARANEAE: DIPLURIDAE) FROM THE LOWER CRETACEOUS CRATO LAGERSTÄTTE, ARARIPE BASIN, NORTH-EAST BRAZIL

Abstract:  The first mygalomorph spiders from the Lower Cretaceous Crato Lagerstätte of Cearà Province, north-east Brazil, are described, from adult males and females, in two new genera and species: Cretadiplura ceara Selden, gen. et sp. nov. and Dinodiplura ambulacra Selden, gen. et sp. nov. They belong to the extant family Dipluridae, hitherto known as fossils only from Tertiary strata; thus this occurrence extends the family record by some 90 myr.     

Morphology of Cambrian lobopodian eyes from the Chengjiang Lagerstätte and their evolutionary significance

Visual organs are widely distributed throughout the animal kingdom and exhibit a great diversity of morphologies. Compound eyes consisting of numerous visual units (ommatidia) are the oldest preserved visual systems of arthropods, but their origins are obscure and hypothetical models for their evolution have been difficult to test in the absence of unequivocal fossil evidence. Here we reveal the detailed eye structures of well-preserved Early Cambrian lobopodians Luolishania longicruris and Hallucigenia fortis from the Chengjiang Lagerstätte, China. These animals possess a pair of eyes composed of at least two visual units, interpreted as pigment cups. Contrary to previous suggestions that Cambrian lobopodians possessed ocellus-like eyes comparable to those of extant onychophorans, this multi-component structure is more similar to the lateral eyes of arthropods. Morphological comparison and phylogenetic analyses indicate that these lobopodian eyes may represent an early stage in the evolution of the ancestral visual system of euarthropods.     

Morphology of Luolishania longicruris (Lower Cambrian,Chengjiang Lagerstätte,SW China) and the phylogenetic relationships within lobopodians

New material of the lobopodian Luolishania longicruris has been recovered from the Lower Cambrian Chengjiang Lagerstätte, southwest China. The specimens throw new light on several morphological features of the species, including the paired antenniform outgrowths, eyes, head shield, setae and other cuticular projections, as well as the differentiated sclerites, appendages, claws, and lobopod interspaces. L. longicruris shows well developed tagmosis: a distinct head and a trunk divided into two sections. The new data allow a revised comparison with other lobopodians. Miraluolishania haikouensis Liu et al., 2004 is considered to be a junior synonym of L. longicruris Hou and Chen, 1989. Evidence from gut filling and specialized morphological characters indicates that L. longicruris may have had a filter feeding lifestyle. A new cladistic analysis suggests that fossil lobopodians are paraphyletic or even polyphyletic and L. longicruris may be an important representative of the stem lineage leading to arthropods.     

RHYNCHONELLIFORMEAN BRACHIOPODS WITH SOFT-TISSUE PRESERVATION FROM THE EARLY CAMBRIAN CHENGJIANG LAGERSTÄTTE OF SOUTH CHINA

Abstract:  Cosmopolitan kutorginates, the most abundant Early Cambrian rhynchonelliformean brachiopods, became extinct at the end of the Middle Cambrian. Consequently, any information concerning the anatomy of this peculiar lineage of brachiopods has great phylogenetic significance with regard to their extant relatives for analogies with the stem-group clade. Such data have been supplied from fossils of which the soft parts have been preserved in exceptional detail. A new brachiopod, Kutorgina chengjiangensis sp. nov., from the Early Cambrian Chengjiang Lagerstätte of southern China, is described here. It is the first articulated brachiopod species collected from this deposit. The specimens preserve a set of soft-body parts, i.e. lophophore, digestive tract and pedicle, all previously poorly known in almost all Palaeozoic calcareous brachiopod taxa. The lophophore attains an early spirolophe stage, clearly homologous to that in the coeval lingulids. The digestive tract consists of a mouth, oesophagus, swollen stomach, intestine and a terminal anus. The pedicle protruding between the valves is stout and elongate, with annulated lamellae on the surface, and contains a putative cœlomic cavity. K. chengjiangensis displays the characteristics of the stem group of calcareous brachiopods, and illustrates anatomical similarities between Cambrian phosphatic- and calcareous-shelled brachiopods, thus corroborating that the Brachiopoda are a monophyletic group.     

Secular distribution of Burgess-Shale-type preservation

Burgess-Shale-type preservation is defined as a taphonomic pathway involving the exceptional organic preservation of non-mineralizing organisms in fully marine siliciclastic sediments. In the Phanerozoic it occurs widely in Lower and Middle Cambrian sequences but subsequently disappears as a significant taphonomic mode. The hypothesis that this distribution derives solely from a secular increase in the depth of bioturbation is falsified: low bioturbation indices do not prevent the rapid enzymatic degradation of organic structure, nor do they account for the conspicuous absence of comparable preservation during the Vendian. An earlier, Late Riphean (ca. 750–850 Ma), interval of enhanced organic-walled fossil preservation suggests a long-term recurrence in Burgess-Shale-type taphonomy that is independent of metazoan activity. A model based on the potentially powerful anti-enzymatic and/or stabilizing effects of clay minerals on organic molecules is proposed to account for Burgess-Shale-type preservation. Long-term changes in average clay mineralogies and the ocean chemistry that determines their interaction with organic molecules are likely to have induced the pronounced secular distribution of these fossil biotas, while regional variations in tectonism, weathering, etc., explain their non-uniform geographic distribution; the close correlation between exceptional, organic-walled fossil preservation and volcano-genic sedimentation in Tertiary lake deposits provides a compelling analogue. Recognition of a temporal control on Burgess-Shale-type preservation constrains the evolutionary scenarios that can be drawn from such biotas; significantly, neither the initial rate of appearance, nor the ultimate fate of Burgess-Shale-type taxa can be directly assessed. □ Taphonomy, exceptional preservation, organic preservation, fossil Lagerstätten, Burgess Shale, clay mineralogy, clay-organic interactions, secular change, Cambrian, Proterozoic.     

An Odontogriphid from the Upper Permian of Australia

A problematic organism, Bowengriphus perphlegis gen. et sp. nov., is described based on two specimens from the Late Permian Rangal Coal Measures of eastern central Queensland. It displays a double-looped feeding apparatus bearing small conical elements, considered homologous with that of the supposed lophophorate Odontogriphus omalus Conway Morris, 1976, from the Middle Cambrian Burgess Shale of British Columbia, Canada. Odontogriphids are thus interpreted as a group that survived through the Palaeozoic and made the transition from marine to freshwater environments. Recent proposals that odontogriphids are stem-group chordates are not well established.     

贵州下寒武统牛蹄塘生物群中的双孔钱包海绵Crumillospongia biporosa

下寒武统澄江生物群、中寒武统布尔吉斯页岩生物群及下寒武统牛蹄塘生物群均产有Crumillospongiabi-porosa,由于产C.biporosa的牛蹄塘生物群下部层位低于澄江生物群,贵州遵义松林可能是C.biporosa的最低层位及发源地。本文描述了贵州遵义下寒武统牛蹄塘生物群的C.biporosa,为C.biporosa的发生、演化及古地理分布提供了重要的信息和证据。     

BROMALITES FROM THE SOOM SHALE LAGERSTÄTTE (UPPER ORDOVICIAN) OF SOUTH AFRICA: PALAEOECOLOGICAL AND PALAEOBIOLOGICAL IMPLICATIONS

Abstract:  Bromalites from the Soom Shale are allocated to five main categories on the basis of shape, content and internal structure: those containing broken conodont elements; those containing brachiopod fragments; corrugated/spiral forms; coiled coprolites and wrinkled coprolites. It is impossible to allocate specific bromalites to the organisms that formed them, but the occurrence of crushed discinoid valves in several specimens demonstrates that an effective durophagous predator was present in the Soom Shale community. The presence of fragmented elements of conodonts in other specimens provides direct evidence of tiering within the predatory trophic level in the Soom Shale. Conodonts, other agnathan vertebrates, orthoconic cephalopods and eurypterids are all possible contenders for producing most of the bromalites recorded, but there may have been unrecorded large predators in the community.     

Nektaspid arthropods from the Lower Cambrian Emu Bay Shale Lagerstätte,South Australia,with a reassessment of lamellipedian relationships

Abstract: The lower Cambrian Emu Bay Shale on Kangaroo Island, South Australia, contains the only known Cambrian Burgess Shale‐type biota in Australia. Two new lamellipedian arthropods, Emucaris fava gen. et sp. nov. and Kangacaris zhangi gen. et sp. nov., from the Emu Bay Shale Lagerstätte are described as monotypic genera that are resolved cladistically as a monophyletic group that is sister to Naraoiidae + Liwiidae and classified within the Nektaspida as a new family Emucarididae. Shared derived characters of Emucarididae involve a bipartite, elongate hypostome and elongation of the pygidium relative to the cephalic shield and very short thorax. A monophyletic Liwiidae is composed of Liwia and the Ordovician Tariccoia + Soomaspis but excludes Buenaspis, and even the membership of Buenaspis in Nektaspida is contradicted amongst the shortest cladograms. New morphological interpretations favour affinities of Kwanyinaspis with Conciliterga rather than with Aglaspidida, and Phytophilaspis with Petalopleura.     

Benthic primary producers in exceptionally preserved Cambrian biotas of North China

Strata situated within the South China Craton host several key Cambrian macroalgal assemblages, including those of the Chengjiang, Kaili, and Tsinghsutung Lagerstätte. In this report, Cambrian macroalgae are described for the first time from the North China Craton, an area that, during the early Paleozoic, was not contiguous with the South China Craton. The material collected from the Mantou Formation and overlying Zhangxia Formation (Miaolingian Series) in Shandong Province is preserved as carbonaceous compressions and includes abundant simple tubular forms. These fall into two groupings on the basis of size and have characteristics broadly comparable to Fuxianospira gyrata and Sinocylindra yunnanensis, both of which are well-known from the Chengjiang biota and may represent simple siphonous green algae. Additional material includes irregular circular to oval films referable to Morania fragmenta and a tuft-like form superficially similar to Marpolia spissa, taxa with possible cyanophyte affinities described initially on the basis of material from the Burgess Shale but also known from the Cambrian of South China. The material is typical of the “Cambrian Flora” and provides further support for the cosmopolitan distribution of this noncalcified macroalgal flora at low latitude settings during the Cambrian.     

A miniscule optimized visual system in the Lower Cambrian

Simultaneously with the development of animal body plans, probably before the Precambrian, there was an explosive diversification of visual systems. Competition of performance in these visual systems was a critical factor in the evolution of life systems.
Here we analyse the visual system in the lobopod Miraluolishania haikouensis ( Liu et al ., 2004 ) from the Lower Cambrian Chengjiang Lagerstätte, Kunming, China. It consists of a very small eye with a miniscule lens. A physical problem lies in the fact that due to the usual refractive conditions of such a lens, it hardly represents an improvement of the visual quality over the basal pit- or pinhole camera eyes. To develop such a lavish visual system, however, would not have been of any value, if it achieved no more than an equal level or represented even a retrograde step in evolutionary progress. We show how this system may have allowed pattern recognition even under poor light conditions. Optimization of such a tiny eye is costly but is not 'a wasted effort' in evolution. In M. haikouensis ( Liu et al. , 2004 ), an excellently adapted miniscule visual system has become possible.