First record of the brachiopod Lingulella waptaensis with pedicle from the Middle Cambrian Burgess Shale

Pettersson Stolk, S., Holmer, L. E. and Caron, J ‐B. 2010. First record of the brachiopod Lingulella waptaensis with pedicle from the Middle Cambrian Burgess Shale. —Acta Zoologica (Stockholm) 91 : 150–162 The organophosphatic shells of linguloid brachiopods are a common component of normal Cambrian–Ordovician shelly assemblages. Preservation of linguloid soft‐part anatomy, however, is extremely rare, and restricted to a few species in Lower Cambrian Konservat Lagerstätten. Such remarkable occurrences provide unique insights into the biology and ecology of early linguloids that are not available from the study of shells alone. Based on its shells, Lingulella waptaensis Walcott, was originally described in 1924 from the Middle Cambrian Burgess Shale but despite the widespread occurrence of soft‐part preservation associated with fossils from the same levels, no preserved soft parts have been reported. Lingulella waptaensis is restudied herein based on 396 specimens collected by Royal Ontario Museum field parties from the Greater Phyllopod Bed (Walcott Quarry Shale Member, British Columbia). The new specimens, including three with exceptional preservation of the pedicle, were collected in situ in discrete obrution beds. Census counts show that L. waptaensis is rare but recurrent in the Greater Phyllopod Bed, suggesting that this species might have been generalist. The wrinkled pedicle protruded posteriorly between the valves, was composed of a central coelomic space, and was slender and flexible enough to be tightly folded, suggesting a thin chitinous cuticle and underlying muscular layers. The nearly circular shell and the long, slender and highly flexible pedicle suggest that L. waptaensis lived epifaunally, probably attached to the substrate. Vertical cross‐sections of the shells show that L. waptaensis possessed a virgose secondary layer, which has previously only been known from Devonian to Recent members of the Family Lingulidae.     

Epithelial moulds from some Upper Ordovician acrotretide brachiopods of Ireland

Three new genera of acrotretid brachiopods are of general morphological interest in that they display polygonal mosaics on the internal shell surfaces, features which are believed to be moulds of the outer epithelial cells. The mosaic is further noted in a specimen of Eoconulus , a form which may also belong to the Acrotretacea. Such mosaics have not previously been recorded from the Acrotretacea. Polygonal mosaics, epithelial moulds, outer epithelium , Eoconulus, Acrotretacea, Ordovician.     

Larval ecology and morphology in fossil gastropods

The shell of marine gastropods conserves and reflects early ontogeny, including embryonic and larval stages, to a high degree when compared with other marine invertebrates. Planktotrophic larval development is indicated by a small embryonic shell (size is also related to systematic placement) with little yolk followed by a multiwhorled shell formed by a free‐swimming veliger larva. Basal gastropod clades (e.g. Vetigastropoda) lack planktotrophic larval development. The great majority of Late Palaeozoic and Mesozoic ‘derived’ marine gastropods (Neritimorpha, Caenogastropoda and Heterobranchia) with known protoconch had planktotrophic larval development. Dimensions of internal moulds of protoconchs suggest that planktotrophic larval development was largely absent in the Cambrian and evolved at the Cambrian–Ordovician transition, mainly due to increasing benthic predation. The evolution of planktotrophic larval development offered advantages and opportunities such as more effective dispersal, enhanced gene flow between populations and prevention of inbreeding. Early gastropod larval shells were openly coiled and weakly sculptured. During the Mid‐ and Late Palaeozoic, modern tightly coiled larval shells (commonly with strong sculpture) evolved due to increasing predation pressure in the plankton. The presence of numerous Late Palaeozoic and Triassic gastropod species with planktotrophic larval development suggests sufficient primary production although direct evidence for phytoplankton is scarce in this period. Contrary to previous suggestions, it seems unlikely that the end‐Permian mass extinction selected against species with planktotrophic larval development. The molluscan classes with highest species diversity (Gastropoda and Bivalvia) are those which may have planktotrophic larval development. Extremely high diversity in such groups as Caenogastropoda or eulamellibranch bivalves is the result of high phylogenetic activity and is associated with the presence of planktotrophic veliger larvae in many members of these groups, although causality has not been shown yet. A new gastropod species and genus, Anachronistella peterwagneri, is described from the Late Triassic Cassian Formation; it is the first known Triassic gastropod with an openly coiled larval shell.     

Recent freshwater oscillatoriacean analogue of the Lower Palaeozoic calcareous alga Angulocellularia

Recent freshwater cyanophyte oncoids from Canandaigua Lake, New York, consist almost entirely of minute irregular, bush-like, micritic masses comparable with the calcareous alga Angulocellularia Vologdin from Lower Cambrian algal-arehaeocyathan bioherms and biostromes of the Siberian Platform and western Mongolia. The Recent specimens arc radially orientated within the oncoids and occasionally contain moulds of axial filaments 15 μm in diameter. Acid-residues of the oncoids yield abundant Schizothrix calcicola. The micritic bush-like masses are interpreted to have formed by the calcification of the sheaths of these cyanophytes. This analogue allows Angulocellularia to be reinterpreted as a calcified oscillatoriacean cyanophyte, rather than a rhodophyte as previously suggested. It indicates that filamentous cyanophytes can produce apparently solid calcareous fossils, not only porostromatc tubes. The occurrence of Angulocellularia has been overlooked in some previous studies of Cambrian and Ordovician bioherms. The generic diagnosis is emended.     

An early Ordovician patelliform gastropod, Palaelophacmaea, reinterpreted as a coelenterate

Yochelson, Ellis L. & Stanley, George D., Jr. 1981 12 15: An early Ordovician patelliform gastropod, Palaelophacmaea , reinterpreted as a coelenterate. Lethaia , Vol. 15, pp. 323–330. Oslo. ISSN W24–1164.
The fossil Palaelophacmaea criola Donaldson, from the early Ordovician Stonehenge Formation of central Pennsylvania, was described as a patelliform gastropod. A reinterpretation of the type lot and study of a few additional specimens provide the basis for an alternative placement. Palaelophacmaea is here assigned to the Hydrozoa, as a possible chondrophore. It has an exceptionally thin shell or test and concentric but irregular corrugations. Cambrian univalve genera having a more or less circular outline that are currently assigned to the Gastropoda or Monoplacophora should be reexamined to see whether they have the features of fossil chondrophore coelenterates rather than those of molluscs. The late Cambrian Palaeoacmaea Hall & Whitfield is removed from the monoplacophoran Mollusca and left unassigned as to phylum. We judge that at least some early Cambrian species of Scenella are probably coelenterate remains. * Early Ordovician , Palaelophacmaea, Gastropoda, Monoplacophora, Hydrozoa .     

The problematic Hadimopanella, Kaimenella, Milaculum and Utahphospha identified as sclerites of Palaeoscolecida

Hinz, I., Kraft, P., Mergl, M. & Müller, K. J. 1990 04 15: The problematic Hadimopanella, Kaimenella, Milaculum and Utahphospha identified as sclerites of Palaeoscolecida. Lefhaia , Vol. 23 , pp. 217–221. Oslo. ISSN 024–1164.
Hadimopanella Gedik, 1 977 , Kaimenella Marss, 1988 and Milaculum Müller, 1973 have been established on the basis of isolated elements of unknown origin. Recently, the latter genus has been tentatively related to the Agnatha (van den Boogaard 1988). By contrast, Bendix-Almgreen & Peel (1988) assigned Hadimopanella to the chordate stock but definitely excluded it from vertebrates. Well-preserved worm-like organisms of Palaeoscolecida Conway Morris & Robison. 1986 are known from the Lower Cambrian to the Lower Ordovician. They have their outer surface covered with a pattern of sequin-like sclerites which evidence the systematic affiliation with the isolated sclerites mentioned above. Based on similar structures on the outer surface, Utahphospha Miiller & Miller, 1976 is considered to belong to the same group. * Early Palaeozoic fossils, Palaeoscolecida, isolated sclerites, phosphatization .     

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.     

GENERIC HOMES FOR BRITISH SILURIAN LINGULOID BRACHIOPODS

Abstract: Since the brachiopod Order Lingulida has been greatly revised in recent years, with particular emphasis on Cambrian and Ordovician genera, many well‐established Silurian linguloid species from the British Isles have been left without an appropriate genus in which to place them. This is rectified here by allocating the various species, mostly erected in the nineteenth or early twentieth centuries, to a variety of genera which have been mainly erected within the last forty years. Two new genera are erected, Mergliella, with type species Lingula bechei Davidson, 1866 , and Striatilingula, with type species Lingula? striata J. de C. Sowerby, 1839 .     

Early Palaeozoic diversification of chitons (Polyplacophora, Mollusca) based on new data from the Silurian of Gotland, Sweden

Isolated, well-preserved silicified sclerites of an unusually diverse Silurian paleoloricate assemblage from Gotland preserve morphological features that are important in interpreting palaeobiology. The typically granular dorsal ornament is comparable with Recent chitons, and is hence possibly linked functionally with sensory aesthetes. Ventral structures particularly in thickened shells indicate major muscle attachment sites sub-apically or, equivalently, beneath the rim of the apical area, and also marginally. In early chitons such as Cambrian Matthevia, deep ventral cavities represent comparable sub-apical sites. Three Gotland genera with an unusual, holoperipheral shell growth style apparently represent plated aplacophorans (cf. Acaenoplax), which coexisted with paleoloricate chitons in shallow inshore carbonate shelf environments. Sclerite features of all the Gotland genera are discussed together since they share most characteristics. The new family Heloplacidae includes those genera and Acaenoplax, which in a preliminary cladistic analysis form a sister group to other Lower Palaeozoic paleoloricates. Multiplated skeletons in paleoloricates and this group of aplacophorans represent parallel evolution of dorsal armour, which in chitons resulted in overlapping, articulating sclerites. The diversity of the mid-Silurian Gotland assemblage is examined against early evolutionary diversification of polyplacophorans, aplacophorans, and in relation to the overall record of Palaeozoic paleoloricate and neoloricate chitons. Peaks in diversity in early Ordovician, mid-Silurian and early Carboniferous times correspond to periods with widespread development of low latitude carbonate shelves. Neoloricates, apparently with an additional shell layer that contributed articulatory plates, appeared in the Devonian from where the fossil record remains poorly known.     

Epithelial cell moulds in acrotretoid brachiopods

The preservation of polygonal imprints of epithelial cells in acrotretoid brachiopods is reviewed and supplemented by new data from the Cambrian of southern Great Britain. The imprints are confirmed as representing moulds of epithelial cells rather than an artefact of microstructure or preserved soft tissues, as they are (1) recorded in most taxa reviewed, (2) best preserved in areas where the shell has been thickened and (3) similar in size to cells recorded in Lingula, the closest living relative to the now extinct acrotretoids. Analysis of the morphology and sizes of epithelial cell moulds demonstrates that there is no consistent relationship between cell width and valve size, and that epithelial cells are not a useful taxonomic character within this group.     

Cambrian biostratigraphy of the Bowers back-arc basin,Northern Victoria Land,Antarctica — A review

The Bowers Mountains in Northern Victoria Land contain the richest Cambrian Series 3 (Miaolingian, middle Cambrian) and Cambrian Series 4 (Furongian, late Cambrian) fossiliferous successions in Antarctica. Almost all the fossils are found within the Bowers Supergroup, which outcrops within the Bowers Terrane, a fault-bounded northwest-southeast oriented strip in Northern Victoria Land. The fossils provide the main age control on the history and evolution of the Bowers volcanic arc and back-arc basin. The great bulk of the fossils occur within the Spurs Formation. The fossil assemblages are dominated by agnostoids and polymerid trilobites with most ranging in age from Drumian to Paibian, although one fauna is of Jiangshanian age. Over 40 agnostoid taxa and over 100 polymerid trilobite taxa have been recorded from the rocks of the Bowers Supergroup. The youngest fauna occurs within the adjacent Robertson Bay Terrane, where a limited fauna of polymerid trilobites and conodonts from within a limestone olistolith have a very late Cambrian or early Ordovician age. Faunal affinities are mainly with Australia, New Zealand, North and South China and the Himalaya with lesser ties to Iran, Kazakhstan, Siberia and Laurentia.     

Muscle scars, shell form and torsion in Cambrian and Ordovician univalved molluscs

Runnegar, Bruce 1981 12 15: Muscle scars, shell form and torsion in Cambrian and Ordovician univalved molluscs. Lethaia, Vol. 14, pp. 311–322. Oslo. ISSN 0024–1164. Well preserved muscle scars have been discovered on one or more specimens of the early Palaeozoic univalved molluscs Pelagiella, Matherella, Sinuites and ?Bucania. From this information it is argued that the bellerophonts Sinuites and ?Bucania were untorted, that Pelagiella had undergone about 10° of torsion, and that Matherella may have been a fully torted, hyperstrophic descendant of Pelagiella. Other early gastropods such as Aldanella could have been independently derived from Pelagiella and therefore the Class Gastropoda may be at least diphyletic. In general, small dextrally coiled archaeogastropods such as Aldanella and the pelagiellids are the most common asymmetrical univalves of the Early Cambrian, but rare ultradextral (hyperstrophic) forms are also known. Only pelagiellids are known from the Middle Cambrian, and ultradextral forms dominate in the Late Cambrian. As a result, slit-bearing archaeogastropods are not common before the Ordovician. Yuwenia bentleyi gen. et sp. nov. is a new taxon proposed in this work. *Mollusca, Gastropoda, Monoplacophora, anatomy, evolution, Cambrian, Ordovician, Yuwenia n.g.     

Cephalopod origin and evolution: A congruent picture emerging from fossils, development and molecules: Extant cephalopods are younger than previously realised and were under major selection to become agile, shell-less predators

Cephalopods are extraordinary molluscs equipped with vertebrate‐like intelligence and a unique buoyancy system for locomotion. A growing body of evidence from the fossil record, embryology and Bayesian molecular divergence estimations provides a comprehensive picture of their origins and evolution. Cephalopods evolved during the Cambrian (～530 Ma) from a monoplacophoran‐like mollusc in which the conical, external shell was modified into a chambered buoyancy apparatus. During the mid‐Palaeozoic (～416 Ma) cephalopods diverged into nautiloids and the presently dominant coleoids. Coleoids (i.e. squids, cuttlefish and octopods) internalised their shells and, in the late Palaeozoic (～276 Ma), diverged into Vampyropoda and the Decabrachia. This shell internalisation appears to be a unique evolutionary event. In contrast, the loss of a mineralised shell has occurred several times in distinct coleoid lineages. The general tendency of shell reduction reflects a trend towards active modes of life and much more complex behaviour.     

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.     

Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny

Popov, L.E., Bassett, M.G., Holmer, L.E., Skovsted, C.B. & Zuykov, M.A. 2010: Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny. Lethaia, Vol. 43, pp. 323–333. Well preserved specimens of the Early Palaeozoic craniiform brachiopods Orthisocrania and Craniops retain clear evidence of a lecithotrophic larval stage, indicating the loss of planktotrophy early in their phylogeny. The size of the earliest mineralized dorsal shell was <100 μm across, and the well preserved shell structure in these fossil craniiforms allows their earliest ontogeny to be compared directly with that of living Novocrania, in which the first mineralized dorsal shell (metamorphic shell) is secreted only after settlement of the lecithotrophic larvae. Immediately outside this earliest shell (early post‐metamorphic or brephic shell) and in the rest of the dorsal valve the primary layer in both fossil and living craniiforms has characteristic radially arranged laths, which are invariably lacking in the earliest dorsal shell. The ventral valve of the fossil specimens commonly preserves traces of an early attachment scar (cicatrix), which is equal in size to the dorsal metamorphic shell, and the brephic post‐metamorphic ventral valve also has a primary shell with radially arranged laths. However, a primary shell with radial laths is completely lacking in the ventral valve of living Novocrania, indicating that heterochrony may have been involved in the origin of the encrusting mode of life in living craniids; the entire ventral valve of Recent craniids (with the possible exception of Neoancistrocrania) may correspond to the earliest attachment scar of some fossil taxa such as Orthisocrania. It is also probable that the unique absence of an inner mantle lobe as well as the absence of lobate cells in Novocrania could be the result of heterochronic changes. The dorsal valve of both fossil and living craniiforms has a marked outer growth ring, around 500 μm across, marking the transition to the adult, and a significant change in regime of shell secretion. The earliest craniiform attachment is considered to be homologous to the unique attachment structures described recently in polytoechioids (e.g. Antigonambonites) and other members of the strophomenate clade. However, unlike the craniiforms, polytoechioids and strophomenates all have planktotrophic larvae, and planktotrophy is most probably a plesiomorphic character for all Brachiopoda. □Brachiopoda, Craniiformea, Early Palaeozoic, ontogeny, phylogeny.     

The origin of annelids

Annelids are a phylum of segmented bilaterian animals that have become important components of ecosystems spanning terrestrial realms to the deep sea. Annelids are remarkably diverse, possessing high taxonomic diversity and exceptional morphological disparity, and have evolved numerous feeding strategies and ecologies. Their interrelationships and evolution have been the source of much controversy over the past century with the composition of the annelid crown group, the relationship of major groups and the body plan of the ancestral annelid having undergone major recent revisions. There is a convincing body of molecular evidence that polychaetes form a paraphyletic grade and that clitellates are derived polychaetes. The earliest stem group annelids from Cambrian Lagerstätten are errant, epibenthic polychaetes, confirming that biramous parapodia, head appendages and diverse, simple chaetae are primitive for annelids. Current evidence from molecular clocks and the fossil record suggest that crown group annelids are a Late Cambrian – Ordovician radiation, with clitellates radiating in the Late Palaeozoic. Their body fossil record is largely confined to deposits showing exceptional preservation and is punctuated by the acquisition of hard parts in major groups. The discovery of an Ordovician fossil with soft tissues has shown that machaeridians are in fact a clade of crown polychaetes. They were in existence for more than 200 million years and possess unique calcitic dorsal armour, allowing their mode of life and phylogeny to be interpreted in the context of the annelid body plan. We identify a novel clade of machaeridians, the Cuniculepadida, which exhibit a series of adaptations for burrowing.     

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...     

The Ordovician Biodiversification: revolution in the oceanic trophic chain

The Early Palaeozoic phytoplankton (acritarch) radiation paralleled a long-term increase in sea level between the Early Cambrian and the Late Ordovician. In the Late Cambrian, after the SPICE δ¹³C_carb excursion, acritarchs underwent a major change in morphological disparity and their taxonomical diversity increased to reach highest values during the Middle Ordovician (Darriwilian). This highest phytoplankton diversity of the Palaeozoic was possibly the result of palaeogeography (greatest continental dispersal) and major orogenic and volcanic activity, which provided maximum ecospace and large amounts of nutrients. With its warm climate and high atmospheric CO₂ levels, the Ordovician was similar to the Cretaceous: a period when phytoplankton diversity was at its maximum during the Mesozoic. With increased phytoplankton availability in the Late Cambrian and Ordovician a radiation of zooplanktonic organisms took place at the same time as a major diversification of suspension feeders. In addition, planktotrophy originated in invertebrate larvae during the Late Cambrian–Early Ordovician. These important changes in the trophic chain can be considered as a major palaeoecological revolution (part of the rise of the Palaeozoic Evolutionary Fauna of Sepkoski). There is now sufficient evidence that this trophic chain revolution was related to the diversification of the phytoplankton, of which the organic-walled fraction is partly preserved.     

THE EFFECTS OF SAMPLING BIAS ON PALAEOZOIC FAUNAS AND IMPLICATIONS FOR MACROEVOLUTIONARY STUDIES

Abstract:  Trilobites, a dominant component of marine faunas during the Cambrian and Ordovician and which survived until the end of the Permian (542–251 Ma) have been used in many macroevolutionary analyses. Here, we use a discovery curve to document the sampling history of trilobites, which we consider a proxy for Palaeozoic faunas in general. At higher taxonomic ranks, orders, suborders and superfamilies, the fossil record has been completely sampled, while the family rank also shows a high level of sampling completeness, having reached an asymptote in 1970. Importantly, this levelling-off occurred even though worker effort continued to increase. However, at genus level the sampling record is incomplete, indicating that families should not be used as a proxy for genera. There is little variation among the different subsets of generic data, with the sampling history of different stratigraphic periods and among different orders being very similar. However, there is noticeable variation among geographical regions, caused by variations in worker effort, and this could cause problems when comparing speciation and diversity patterns across faunal provinces. The role of synonyms on sampling history has had little effect.