A Stem Group Brachiopod From The Lower Cambrian: Support For A Micrina (Halkieriid) Ancestry

The shell structure of the Lower Cambrian Mickwitzia , a bilaterally symmetrical bivalve hitherto doubtfully assigned to the Brachiopoda, confirms that the genus shares characters with linguliform brachiopods. The columnar lamination of its organophosphatic shell is homologous with that characterizing acrotretides. The shell, however, is also pervaded by striated apatitic tubes indistinguishable from those permeating the sclerites of the problematic organophosphatic, laminar–shelled Micrina which is close to Halkieria . No crown group brachiopods have such tubes that are presumed to have contained setae. The presence of both these features in the Mickwitzia shell suggests that the stock is a stem group brachiopod with a halkieriid ancestry.     

Epibionts on the lingulate brachiopod Diandongia from the Early Cambrian Chengjiang Lagerst?tte,South China

The classic Chengjiang Lagerstätte (Lower Cambrian, Atdabanian stage: Yu''anshan Formation) Yunnan, southwestern China, has yielded, besides the exceptional and often controversial soft-bodied fossils, a fauna of primitive/early lingulid brachiopods. Diandongia pista (Rong 1974) is one of the commonest and most strongly mineralized of the phosphatic brachiopods from the Lagerstätte. The shells of this species have been found to commonly serve as a basibiont host. Epibionts comprise the coeval brachiopod Longtancunella chengjiangensis and the cone-shaped cnidarian-related Archotuba conoidalis, as well as rounded smaller-sized epizoans (lesser than 2 mm). A principle morphological analysis demonstrates that the ovoid and rounded organisms that often occur along the commissure of D. pista resemble small juvenile or immature brachiopods. Epibiont-bearing shells of D. pista with soft-tissue preservation demonstrate that the host brachiopods were overgrown while alive, and provide an argument for D. pista having a semi-infaunal life style with only the slim pedicle embedded in sediment. The epibiotic association sheds direct light on the ecology of Cambrian brachiopods in soft-substrate marine environments. The Chengjiang fossils demonstrate that the Early Cambrian brachiopods, as compared with recent lingulids, occupied different and a wider spectrum of ecological niches and tiers of space.     

A new family of Cambrian rhynchonelliformean brachiopods (Order Naukatida) with an aberrant coral‐like morphology

Tomteluva perturbata gen. et sp. nov. and Nasakia thulensis gen. et sp. nov., two new rhynchonelliformean brachiopod taxa, are described from carbonate beds from the lower middle Cambrian (Series 3, Stage 5) basinal Stephen Formation, Canada, and the upper lower Cambrian (Series 2, Stage 4) Henson Gletscher Formation, North Greenland, respectively. The two taxa are characterized by an unusual coral‐like morphology typified by a high conical ventral valve with an anteriorly curved umbo and a tube‐like structure inside the ventral valve, interpreted as pedicle tube. Both resemble the problematic late middle Cambrian (Drumian) species Anomalocalyx cawoodi Brock from Australia, whose systematic affiliation is controversial. Together, the three genera are interpreted as representatives of a new family of rhynchonelliformean brachiopods, the Tomteluvidae fam. nov., which is interpreted as an aberrant or derived taxon within the Order Naukatida. Convergence between the Tomteluvidae and the coralla of small solitary Cambrian coralimorphs, as well as the late Palaeozoic reef‐building richthofenioid brachiopods, might indicate adaptation to a similar life habits and environments. However, their small size (length 4 mm), well‐developed pedicle and perfect morphological symmetry make it more likely that tomteluvids lived attached to frondose algae or sponges, above the seafloor, in a similar fashion to the acrotretoid brachiopods with which they show a high degree of morphological convergence. Morphological features of the pedicle tube of N. thulensis suggest that the tomteluvid pedicle is homologous to that in modern rhynchonelliformean brachiopods. This is the first evidence of the pedicle type within the Naukatida and represents the oldest confirmation of a rhynchonellate pedicle.     

Cavity-dwelling organisms in Lower Cambrian patch reefs from southern Labrador

An intact and well-preserved biota of cavity-dwelling organisms (coelobionts) of early Cambrian age is found within growth-framework cavities in archaeocyathid patch reefs of the lower Forteau Formation (upper Bonnia-Olenellus Zone) of southern Labrador. The biota was diverse, and consisted of encrusting and vagrant organisms. Renalcis , a Renalcis-like form, Epiphyton, Girvanella , and Serligia , all lived attached to walls and roofs of the cavities; other attached forms were Bija, Archaeotrypa , and two types of Wetheredella , an agglutinated foraminifer. Other organisms, including trilobites, probable calcareous ostracodes, brachiopods, echinoderms, and worms, as well as fungi, are also found preserved within the cavities. The structure of the coelobiontic community is surprisingly modern; it indicates that as early as the Lower Cambrian coelobiontic organisms were well-established in reefs, and were an important part of the reef community as a whole.     

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.     

Evidence of lophophore diversity in early Cambrian brachiopoda

The lophophore, an essential organ of the Brachiopoda, has been used widely in evolutionary and advanced phylogenetic studies, but is hitherto unknown in the fossil record. Here, the extraordinarily well-preserved lophophores of two inarticulated brachiopods Lingulella chengjiangensis and Heliomedusa orienta, from the Lower Cambrian Chengjiang fauna (Yunnan, China) are described. These primitive lophophores, respectively, trocholophous and schizolophous, have some key characters that may be plesiomorphies inherited by their recent descendants. This discovery provides direct evidence regarding the taxonomy, ecosystems and early evolution of inarticulated brachiopods.     

The monophyletic origin of the Brachiopoda

Although it is commonly acepted that the brachiopods descended from phoronid-like ancestors there is dispute over their origin. Traditionally they have been regarded as a monophyletic group, a clade. More recently it has been claimed that brachiopods are polyphyletic and that several of the orders arose independently from separate phoronid-like stocks. The latter point of view implies that brachiopods are not a taxon but merely a grade of organization. Traditional stratophenetic approaches do little to resolve the problem, which may be outside their domain. It is possible, even probable, that the initial radiation involved organisms that lacked mineralized shells. Cladistic analysis of both living forms and Lower Paleozoic taxa strongly supports the contention that brachiopods are monophytetic and closely related to the phoronids. It suggests, however, that the 'inarticulate' Paterinida and Kutorginida are genealogically more closely related to the Articulata than they are to the remaining Inarticulata. □ Brachiopoda, Lophophorata, cladistics, Cambrian.     

Morphology,taxonomic position,and stratigraphic distribution of the early Cambrian skeletal problematics <Emphasis Type="Italic">Mobergella radiolata</Emphasis> Bengtson, 1968

Morphology of the phosphatic sclerites of the species Mobergella radiolata that come from the Lower Cambrian sections is studied, and the stratigraphic distribution of the species is revised. The sclerite laminated microstructure of this species is described and is shown to be similar to that of some gastropods, bivalves, brachiopods, bryozoans, and corals. The species M. hexactina Skovsted, 2003 and M. sibirica Skovsted, 2003 are shown to be junior synonyms of M. radiolata based on sclerite morphology supported by the results of statistical analysis.     

Microstructure of the linguliformean brachiopod Linnarssonia from the Lower Cambrian of Sichuan, China

Acrotretoids, one of the oldest brachiopod groups, are abundant in the Lower Cambrian Jiulaodong Formation. The shell of Linnarssonia sp. is composed of two layers: a primary layer and a columnar secondary layer. The primary layer is mostly exfoliated, resulting in exposure of the openings to the central canal of the columns. Filae are seen on the surface of the columnar layer, indicating that the columnar secondary layer has influenced changes in ornament on the shell surface. The larval shell has only very weak ripples; the post-larval shell has obvious concentric ribs. Small pits of variable shape cover almost the entire shell surface. The secondary layer is composed of several columnar laminations, each of which comprises both the upper and lower laminae and the cylindrical columns between them. On the inner side of shell the thin columnar laminations increase. The new microstructural data show that two shell layers are developed in Early Cambrian acrotretoid brachiopods; the columnar lamination may be a primitive feature of the microstructural development of the Brachiopoda and may help establish the affinity between different stem-group brachiopods.     

A developmental basis for the Cambrian radiation

As extant bilaterian phyla emerged during the Lower Cambrian, these clades acquired morphological features that separated them from their stem groups. At the same time, morphological variants on the body plan within a phylum emerged that we recognize as classes and subphyla. In many cases, the emergence of body plan variants within a phylum is associated with major changes in patterns of early regional specification. Subsequently these different patterns of regional specification were stabilized, probably because later developmental events depended on them. As a consequence, the frequency of new body plan variants involving early development declined in these lineages at later periods during their history. This hypothesis is explicated here by examining the process of early regional specification in the different subphyla of brachiopods and in pairs of species from the same subphylum belonging to different clades that originated during the Jurassic, Ordovician, and Lower Cambrian.     

山东潍坊寒武系苗岭统乌溜阶馒头组上部的腕足动物

寒武系腕足动物属种多样性高、个体数量丰富、形态差异明显、地理分布广泛,具有辅助寒武系三叶虫生物地层划分和对比的潜力.华北板块寒武系苗岭统沉积和化石记录发育良好,是中国苗岭统的经典研究区之一.前人己针对华北寒武系苗岭统乌溜阶腕足动物的系统古生物学开展了一系列基础工作,但这些相关研究主要集中于辽宁地区,目前对华北其他地区苗...     

Boring Late Cambrian organisms

Specimens of inarticulate brachiopods (family Acrotrctidae) with boreholes were found in Upper Cambrian carbonates in the southern Great Basin of the United States. Some morphologic features and preferred orientation of the borings are similar to those made by predators and previously reported in the fossil and Holocene record. Such predatory activity on brachiopods is previously unknown in Cambrian rocks. Taxa associated with these specimens are not known to have been predators and identity of the predatory organism is unknown. Cambrian brachiopods, predation, paleoecology.     

A new family of inarticulate brachiopods from the Lower Cambrian of northwestern Mongolia

A new genus of calcareous-shelled brachiopods, Egyngolella, with E. lenae sp. nov. selected as the type species, is described from the upper part of the Egiin Gol Formation, Atdabanian Stage, Lower Cambrian of northwestern Mongolia and is assigned to a new family that is tentatively placed in the order Obolellida. E. lenae sp. nov. is characterized by an unusual combination of features typical for different, sometimes widely divergent brachiopod lineages.     

Origin and dispersal of the earliest brachiopods

Brachiopods first appeared at the very beginning of the Phanerozoic together with the first skeletal organisms. Most brachiopod taxa that arose in the first half of the Cambrian had a short temporal range and became completely extinct by the middle of the Middle Cambrian. Rigid articulation of the valves of brachiopods was provided by various structures, which also appeared in the Early Cambrian. This fact points to the importance of this feature for the formation of the whole group and at the same time testifies to the high variability of rigid articulation at the early stages of brachiopod evolution. This is a typical manifestation of archaic diversity in this animal phylum, which appeared very early in the Phanerozoic. Another important property of the archaic diversity of the early brachiopods was the large number of centers of diversification. As for the majority of groups, climatic zonality was the main factor determining the distribution of brachiopods at the beginning of the Phanerozoic. The main ecological types of brachiopods also appeared in the Early Cambrian.     

AN ORDOVICIAN LOBOPODIAN FROM THE SOOM SHALE LAGERSTÄTTE, SOUTH AFRICA

Abstract:  The first lobopodian known from the Ordovician is described from the Soom Shale Lagerstätte, South Africa. The organism shows features homologous to Palaeozoic marine lobopodians described from the Middle Cambrian Burgess Shale, the Lower Cambrian Chengjiang biota, the Lower Cambrian Sirius Passet Lagerstätte and the Lower Cambrian of the Baltic. The discovery provides a link between marine Cambrian lobopodians and younger forms from the Silurian and Carboniferous. The new fossil preserves an annulated trunk, lobopods with clear annulations, and curved claws. It represents a rare record of a benthic organism from the Soom Shale, and demonstrates intermittent water oxygenation during the deposition of the unit.     

Traces of predation in the Cambrian

Series two marks a revolution in Cambrian predation when new predators and new predation methods appeared, which led to general increase in predation intensities and in the diversity of prey groups. The number of bored taxa and taxa with the predation scars is similar in the Cambrian. Most of the borings are associated with brachiopods and most of the scars with trilobites. Brachiopods, arthropods, molluscs, cnidarians and echinoderms were the most common prey in the Cambrian. The Cambrian record of predation is dominated by damage inflicted on brachiopods and trilobites. The fossils with predation signs are known from a majority of paleocontinents and all the Cambrian series.     

Early Cambrian coelobiontic communities in tectonically unstable crevices developed in Neoproterozoic andesites, Ossa-Morena, southern Spain

A well-preserved biota of Lower Cambrian cavity-dwelling organisms is recorded within fissures in Neoproterozoic andesites in Ossa-Morena (southern Spain). The cavities are unique among described Lower Cambrian coelobiontic communities due to the igneous character of the host rock. Coelobiontic habitat was episodically enlarged by synsedimentary tectonic fracturing reflecting polyphase infill of recurrent facies. The pioneer coelobiontic biota was diverse, and consisted of encrusting stromatolites and thromboids (dominated by Epiphyton and Renalcis), attached to walls and ceilings of the cavities, associated with archaeocyaths. Sponge spicules and chancelloriid sclerites occur as dense clusters indicating in situ growth, death and decay of spiculate sponges and coeloscleritophorans. Other organisms, such as echinoderms, trilobites and brachiopods, are also found within the cavities as reworked skeletons, indicating that they were washed in from the overlying, open seafloor. The main feature of the coelobiontic biota is the dominance of a sessile, chemosynthetic and filter-feeding epibenthos, composed of microbial communities, archaeocyaths, spiculate sponges (demosponges and rarer hexactinellides) and coeloscleritophorans.     

First record of a bivalved larval shell in Early Cambrian tommotiids and its phylogenetic significance

Abstract: Brachiopods are marine Lophotrochozoa whose soft parts are enclosed in a bivalved shell. Although brachiopods are represented by a rich record from the Early Cambrian to the present, the origin of their bivalved body plan remains controversial. The Early Cambrian organophosphatic tommotiids Micrina and Paterimitra from Australia have been proposed as stem brachiopods. Here, we describe their earliest ontogeny, indicating that tommotiids possessed bivalved planktotrophic larvae. The curious combinations of characters in Micrina and Paterimitra indicate that they may belong to the stems of the Linguliformea and Rhynchonelliformea, respectively. The bivalved shell of adult living brachiopods may represent a plesiomorphic character retained from planktic tommotiid larvae; the crown group body plan of the Brachiopoda may have evolved through the paedomorphic retention of a bivalved larval state.     

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.