Lars E. Holmer

A stem group brachiopod from the Lower Cambrian - support for a Micrina (Halkieriid) ancestry.

Small shelly fossils are common in sediments of Early Cambrian age and include the earliest common representatives of metazoan animals with mineralized hard parts. The group includes fossils of very different morphology, composition and ultrastructure. They seem to represent skeletal remains of numerous animal groups, the biological affinities of which are largely unresolved. However, the wide geographic range of many forms has the potential to enhance biostratigraphic and palaeogeographic resolution in the Early Cambrian. The late Early Cambrian sequence of North-East Greenland has yielded an assemblage of more than 88 species of small shelly fossils, brachiopods and trilobites, indicative of a middle Dyeran age (Botoman equivalent). The recovered fossils include a number of species that are known from other Early Cambrian palaeocontinents, and particularly strong ties to late Early Cambrian faunas of Australia are documented. The many cosmopolitan taxa thus identified suggests a close juxtaposition of palaeocontinents at this time. The systematic affinity of many of these small shelly fossils is poorly understood, partly because of their fragmentary nature and poor preservation. However, new data from North-East Greenland improves our understanding of the function and biological affinity of certain taxa. Collections of the problematic fossil Mongolitubulus from North and North-East Greenland exhibit characters indicative of a defensive function as spines of bivalved arthropods, while species of the problematic genus Triplicatella represent the opercula of an unknown tubular shell, probably related to orthothecid hyoliths. The bivalved fossil Mickwitzia from North-East Greenland combines characters of linguliform brachiopods and sclerites of Micrina, a non-bivalved problematic form (halkieriid) from Australia. The combination suggests that Mickwitzia is a stem group brachiopod and strengthens arguments for a halkieriid ancestry of the brachiopod phylum.

The Scleritome of Eccentrotheca from the Lower Cambrian of South Australia : Lophophorate affinities and implications for tommotiid phylogeny

The first partially articulated scleritome of a tommotiid, Eccentrotheca sp., is described from the Lower Cambrian of South Australia. The Eccentrotheca scleritome consists of individual sclerites; fused in a spiral arrangement, forming a tapering tube-shaped skeleton with an inclined apical aperture and a circular to subcircular cross section. Traditionally, tommotiid sclerites have been assumed to form a dorsal armor of imbricating phosphatic plates in slug-like bilaterians, analogous to the calcareous sclerites of halkieriids. The structure of the Eceentrotheca scleritome is here reinterpreted as a tube composed of independent, irregularly shaped sclerites growing by basal-marginal accretion that were successively fused to form a rigid, protective tubular structure. The asymmetrical shape and sometimes acute inclination of the apical aperture suggests that the apical part of the scleritome was cemented to a hard surface via a basal disc, from which it projected vertically. Rather than being a vagrant member of the benthos, Eccentrotheca most likely represented a sessile, vermiform filter feeder. The new data suggest that the affinities of Eccentrotheca, and possibly some other problematic tommotiids, lie with the lophophorates (i.e., the phoronids and brachiopods), a clade that also possesses a phosphatic shell chemistry and a sessile life habit.

Cambrian‐Ordovician lingulate brachiopods from Scandinavia, Kazakhstan, and South Ural Mountains

Lingulate brachiopods are described from the Upper Cambrian - Lower Ordovician (Tremadoc-Arenig) of Scandinavia (Sweden, Denmark, and Norway), South Ural Mountains, northeastern Central Kazakhstan, and the southern Kendyktas Range in southern Kazakhstan. The faunas comprise a total of 56 species of which 20 are new these are assigned to 40 genera, of which the lingulids Agalatassia and Keskentassia, the siphonotretid Siphonotrerella. and the acrotretids Galinella, Longipegma, Ottenbyella, Akmolina, Mamatia, Sasyksoria, and Otariella are new. The new Subfamily Elliptoglossinae is proposed. The poorly known Cambrianardovician stratigraphy of the South Urals. northeastern Central Kazakhstan, and the southern Kendyktas Range is reviewed. Many sequences in these areas that were previously referred to the Upper Cambrian and Tremadoc can now be correlated with the lower Arenig Hunneberg Stage in Baltoscandia. Three main types of faunal assemblages can be distinguished: (1) the Broeggeria assemblage; (2) several microbrachiopod assemblages; and (3) the Leptembolon-Thysanotos assemblage. The Broeggeria assemblage is distributed world-wide in the Tremadoc of the southern Kendyktas Range, Scandinavia, Belgium, Great Britain, Canada, and Argentina, while the Leptembolon-Thysanotos assemblage is confined to the Arenig of an area surrounding the East European platform, including northern Estonia, Poland, Germany, Bohemia, Serbia, and the South Urals. The microbrachiopod assemblages are known mainly from the Upper Cambrian - Arenig of Scandinavia, South Ural Mountains, northeastern Central Kazakhstan, and the southern Kendyktas Range.

Shell structure and inferred growth, functions and affinities of the sclerites of the problematic Micrina

The stratiform laminae of Micrina sclerites originally consisted of rheomorphic successions of monolayers of micrometric–sized, apatitic tablets, presumably interleaved with chitin and glycosaminoglycans (GAGs). Paired laminae enclose slot–like chambers swelling into lobes distally that originally contained GAGs and deposits of spherulitic and prismatic apatite. The laminae are pervaded by apatitic tubes, apparently secreted by microvillous setoblasts and containing, at the surface, chitinous setae. Internal markings suggest that the triangular (sellate) sclerite supported a pair of muscles and the planospiral (mitral) sclerite, a medial muscle and gonadal sacs flanked by a pair of crescentic muscle bases. Both sclerites were secreted by a mantle with a circumferential fold. The sellate and mitral sclerites are homologized with the anterior and posterior shells of Halkieria and could have become the dorsal and ventral valves of the ancestral brachiopod by a sequence of transformations. These include: the folding of the halkieriid body axis; accelerated mixoperipheral growth of the anterior (dorsal) shell to enclose, with the posterior (ventral) shell, a mantle cavity lined with modified ciliated epithelium of the foot; reduction of sclerite–secreting epithelium to the locus of the brachiopod pedicle epithelium; and the anterior (dorsal) spread of gonadal lamellae.

The scleritome of Paterimitra: an Early Cambrian stem group brachiopod from South Australia

Early Cambrian tommotiids are problematic fossil metazoans with external organophosphatic sclerites that have been considered to be basal members of the lophophorate stem group. Tommotiids are almost exclusively known from isolated or rarely fused individual sclerites, which made previous reconstructions of the actual organism highly conjectural. However, the recent discovery of the first articulated specimens of the tommotiid Eccentrotheca revealed a tubular sclerite arrangement (scleritome) that limited the possible life habit to sessile filter feeding and thus further supported a lophophorate affinity. Here, we report the first articulated specimens of a second tommotiid taxon, Paterimitra from the Early Cambrian of the Arrowie Basin, South Australia. Articulated specimens of Paterimitra are composed of two bilaterally symmetrical sclerite types and an unresolved number of small, asymmetrical and irregular crescent-shaped sclerites that attached to the anterior margin of the symmetrical sclerites. Together, the sclerites form an open cone in which the symmetrical sclerites are joined together and form a small posterior opening near the base of the scleritome, while the irregular crescent-shaped sclerites defined a broad anterior opening. The coniform scleritome of Paterimitra is interpreted to have attached to hard substrates via a pedicle that emerged through the small posterior opening (sometimes forming a tube) and was probably a sessile filter feeder. The scleritome of Paterimitra can be derived from the tubular scleritome of Eccentrotheca by modification of basal sclerites and reduction in tube height, and probably represents a more derived member of the brachiopod stem group with the paired symmetrical sclerites possibly homologous to brachiopod valves.

The Early Cambrian tommotiid Micrina ,a sessile bivalved stem group brachiopod

The tannuolinid Micrina belongs to the tommotiids—a common and widely distributed, but poorly understood, group of Early Cambrian fossil metazoans with multiple external organophosphatic sclerites. Recent findings of sessile articulated tommotiid scleritomes indicate that previous reconstructions of tommotiids as slug-like bilaterians with a dorsal cover of sclerites require detailed re-evaluation. Comparative ultrastructural work has already indicated that the tommotiids might be a sister group to the Brachiopoda, with Micrina representing the most derived and brachiopod-like bimembrate tommotiid. Here we further develop and strengthen this controversial phylogenetic model with a new reconstruction of Micrina, where the two types of sclerites—mitral and sellate—belong to a near bilaterally symmetrical bivalved sessile organism. This new scleritome configuration was tested by recreating an articulated bivalved Micrina from isolated mitral and sellate sclerites; both sclerites have muscles that would have enabled movement of the sclerites. The mitral and sellate sclerites of Micrina are considered to be homologous with the ventral and dorsal valves, respectively, of organophosphatic linguliform brachiopods, indicating that a simple type of filter-feeding within an enclosed bivalved shell had started to evolve in derived tannuolinids. The new reconstruction also indicates that the phylogenetic range of ‘bivalved’, sessile lophophorates is larger than previously suspected.

Gondwanan faunal signatures from Early Palaeozoic terranes of Kazakhstan and Central Asia : evidence and tectonic implications

Abstract Two separate tectonic blocks in the southwestern segment of the Kazakhstanian orogen, the Chu–Ili terrane and the Karatau–Naryn terrane (with particular attention to Malyi Karatau), are selected to illustrate their geological history and major biogeographical signatures from the Cambrian to the early Silurian. Mid- to Late Ordovician brachiopod and trilobite faunas of Chu–Ili show increased endemicity of shallow shelf assemblages, whereas distinct links to equatorial (‘east’) peri-Gondwanan are more evident in trilobite assemblages of the outer shelf. In the Late Ordovician, strong biogeographical affinities to equatorial Gondwanan faunas became firmly established and they are also traceable into the Silurian. Early Cambrian faunas of Malyi Karatau show remarkable similarity to those of South China. From the Middle Cambrian this region evolved as an isolated carbonate seamount, but until the Early Ordovician links to South China faunas were still evident. Benthic faunas from both regions show weak links to contemporaneous faunas of Baltica and little in common with Cambrian and Ordovician faunas of the Siberian craton. This suggests their location in low southern latitudes, in relative proximity to East Gondwana, which places some constraints on plate-tectonic reconstructions in relation to the southern cluster of Kazakhstanian terranes, including Karatau–Naryn, North Tien Shan and Chu–Ili.

Brachiopods: Cambrian-Tremadoc precursors to Ordovician radiation events

Abstract Brachiopod-dominated palaeocommunities incorporating a structure typical of faunal groups within the Palaeozoic Evolutionary Fauna were already present in North and East Gondwana and associated terranes as early as the mid-Cambrian, confined exclusively to shallow marine, inshore environments. The late Cambrian and Tremadoc record of these faunas is incomplete, because of pronounced global sea-level lowstand and subsequent break-up and destruction of the Cambrian Gondwanan margin. It is likely, however, that those groups later forming the core of the Palaeozoic Evolutionary Fauna evolved originally in shallow-water environments of low-latitude peri-Gondwana, and dispersed widely when favourable ecological conditions developed. Conspicuous sea-level rise through the early to mid-Arenig provided newly available habitats in the expanding epeiric seas, where the new faunas evolved and diversified by the mid-Ordovician, when rapid drift separated the early Palaeozoic continents. Relatively short-lived precursor and transitional brachiopod assemblages can be identified on most of the main palaeocontinents prior to the Ordovician radiation of the Palaeozoic Evolutionary Fauna.

Organophosphatic brachiopods : Patterns of biodiversification and extinction in the early Palaeozoic

From a database of 226 Cambrian - Ordovician genera of organophosphatic-shelled brachiopods comprisingthe Subphylum Linguliformea, 11 sets of morphological characters typify all orders and superfamilies. Seven sets of these large-scale evolutionary novelties were established already by the end of the mid Cambrian, 2 more by the end of the late Cambrian, and the remaining 2 before the end of the Arenig. The earliest linguliformeans are of Tommotian age and represent some of the oldest known benthic organisms with a mineralised skeleton. Major diversification at the generic level took place during the mid and late Cambrian, by which time members of the Order Lingulida spread from near-shore to deep-water environments and became dominant in low diversity benthic assemblages that inhabited mobile sandy bottoms. There was a significant decline in diversity of linguliformeans during the latest Cambrian. Following recovery in the late Tremadoc-early Arenig, they then became one of the most distinctive components of benthic assemblages inhabiting marginal environments, e.g. eutrophic basins, shallow mobile sands, and abyssal depths. During the Llanvirn, there was a significant, worldwide turnover in linguliformean brachiopod faunas, when the majority of epibenthic lingulides of the families Obolidae, Zhanatellidae and Elkaniidae became extinct and were replaced, in shallow marine biofacies, by assemblages dominated by bivalved molluscs and burrowing lingulides. There is no trace until the mid Ashgill of a significant decline of micromorphic taxa in linguliformean assemblages characteristic of outer shelf environments. However, all siphonotretides, paterinides and most acrotretide genera disappeared towards the end of the late Ordovician pre-Hirnantian Dicellograptus anceps Biozone.

Homologous skeletal secretion in tommotiids and brachiopods

Tommotiids are distinctive components of the early Cambrian small shelly fauna, almost invariably represented by isolated phosphatic sclerites derived from a multielement protective cover (scleritome). The unusual range of tommotiid sclerite morphologies and unknown construction of the scleritome have severely hampered our understanding of their phylogenetic affinities. However, recent description of rare, articulated scleritome material belonging to the tommotiid genera Eccentrotheca and Paterimitra support the hypothesis that some tommotiids fall within the stem group of the lophophorate phyla Phoronida and Brachiopoda and that at least some tommotiid sclerites are homologous precursors of the shells of organophosphatic brachiopods. Here we show that the shell microstructure of Eccentrotheca and Paterimitra share substantial similarities with paterinid brachiopods. While paterinids possess an overall brachiopod morphology, their microstructure appears more similar to Eccentrotheca and Paterimitra than to nonpaterinate lingulids. These findings strongly support the existence of a brachiopod total group that is solidly rooted within tommotiids, and identify the organophosphatic skeletal composition as plesiomorphic with calcareous shells as derived. The microstructural changes of the proposed tommotiid-brachiopod transition probably reflect an adaptation to fluctuating food and phosphorous intake that came with the switch to a sessile life style at the base of the tommotiid clade.