Homologous shell microstructures in Cambrian hyoliths and molluscs

Hyoliths were among the earliest biomineralizing metazoans in Palaeozoic marine environments. They have been known for two centuries and widely assigned to lophotrochozoans. However, their origin and relationships with modern lophotrochozoan clades have been a longstanding palaeontological controversy. Here, we provide broad microstructural data from hyolith conchs and opercula from the lower Cambrian Xinji Formation of North China, including two hyolithid genera and four orthothecid genera as well as unidentified opercula. Results show that most hyolith conchs contain a distinct aragonitic lamellar layer that is composed of foliated aragonite, except in the orthothecid New taxon 1 that has a crossed foliated lamellar microstructure. Opercula are mostly composed of foliated aragonite and occasionally foliated calcite. These blade or lath‐like microstructural fabrics coincide well with biomineralization of Cambrian molluscs rather than lophophorates, as exemplified by the Cambrian members of the tommotiid‐brachiopod linage. Accordingly, we propose that hyoliths and molluscs might have inherited their biomineralized skeletons from a non‐mineralized or weakly mineralized common ancestor rather than as a result of convergence. Consequently, from the view of biomineralization, the homologous shell microstructures in Cambrian hyoliths and molluscs strongly strengthen the phylogenetic links between the two groups.     

Hyolith-type microstructure in a mollusc-like fossil from the Early Cambrian of Yunnan, China

A phosphatized bilaterally symmetrical, spirally coiled conch from the Lower Cambrian phosphates of the Meishucunian Stage of eastern Yunnan, China, is composed of two well-preserved layers. The outer one is built of longitudinal mineralized fibres not strictly parallel to each other, but may anastomose or branch. The fibres form comarginal ribs with transversal grooves between them, corresponding to growth stages. In the ribs, the fibres are arranged closely and are steeply inclined towards the apex, while in the grooves the fibres are subparallel to the conch surface and more loosely packed. Small elliptic and triangular holes are situated within and between the fibres of the outer layer. The inner layer consists of transversal fibres running around the conch. They are almost parallel to each other but partly separated by narrow discontinuous slits. Despite overall morphological similarity of the conch to shells of some early Cambrian molluscs, the observed orthogonal configuration of longitudinally oriented fibrous structures of the outer layer and transversal fibres of the inner layer is found in orthothecid hyoliths, phosphatized microstuctures of which have been described from the Lower Cambrian of the Siberian Platform. The new material from the Yangtze Platform supports a conclusion of a distinctive type of microstructure available in hyoliths different from molluscan microstructures available from the Lower Cambrian.     

A revision of Holm's Early and early Mid Cambrian hyoliths Of Sweden

ABSTRACT. Re-examination of type specimens of Early Cambrian and early Mid Cambrian hyoliths from Sweden confirms placement of Hyolithes teretiusculus Linnarsson in Hexitheca Syssoiev, and reassignment to the order Hyolithida rather than Orthothecida. Inclusion of Hyolithes affinis Holm within Decoritheca Syssoiev is re-confirmed, and Hyolithes socialis Linnarsson is now referred to Nevadotheca Malinky and Slapylites Marek. An operculum included under Hyolithes oelandicus Holm is transferred provisionally to Maxilites Marek, and conchs of that species are placed under Tulenicornus Val'kov with question. No other hyolith species of Holm can be definitely assigned to genus. The rarity of well preserved opercula is a major impediment to recognizing hyolith taxa confidently, and we follow Holm in attributing the comparative rarity of opercula to the fact that in some hyolith taxa the operculum may have been composed of a different, and perhaps less stable, material than the rest of the conch.     

早寒武世梅树村期软舌螺壳孔沟系统的初步研究

对滇东会泽雨碌、寻甸摆宰早寒武世梅树村期软舌螺壳质微细结构的研究表明,软舌螺壳的孔沟系统十分发育,壳的外层由分叉与合并呈纵向排列的长纤束组成,内层由横向分布的粒柬组成。在外层纤柬之间和纤柬内分布有大量的三角形或长椭圆形的孔洞,这些孔洞的长轴平行于壳的生长方向;在内层粒束之间或粒柬内有横向分布的长沟。内外层的沟孔可以相通。在一些软舌螺纵向和横向的切片中,均发现早期磷酸岩化作用中已被磷质颗粒充填的孔沟形态,这些孔沟分布相当密集,基本垂直壳面,几乎通达壳壁的顶底。多孔性的壳既反映了软舌螺对其特殊的生理和生态功能的要求,也反映了早期带壳动物的原始性和它们在早期生物矿化作用中的特殊地位。     

华北寒武系第二统辛集组软舌螺动物化石群*

软舌螺动物是华北板块寒武系小壳化石基本组成之一,也是寒武纪生命大爆发在华北地区的重要见证者。本文系统描述了华北板块南缘-西南缘(河南-陕西地区)寒武系第二统辛集组软舌螺化石9属14种及1未定种。包括直管螺类6属9种:Conotheca australiensis、Cupitheca holocyclata、Cupitheca costellata、Cupitheca? decollata、Tegminites hymenodes、Triplicatella disdoma、Triplicatella xinjia、Protomicrocornus triplicensis和Longxiantheca mira;软舌螺类3属5种及1未定种:Microcornus eximius、Microcornus petilus、Parkula cf. esmeraldina、Parkula bounites、Parakorilithes mammillatus和Parakorilithes sp.。基于大量保存的精美聚合标本,本文准确重建了锥壳和口盖在软舌螺不同属种中的配置。此外,新材料显示华北辛...     

Biology of the Hyolitha

Hyoliths are Paleozoic fossils that have a calcareous exoskeleton consisting of an elongate, usually bilaterally symmetrical cone, a close fitting operculum, and a pair of curved appendages. Their skeletal ultrastructure resembles the crossed-lamellar shell layers of some molluscs. Several specimens from the Ordovician of France and the Cambrian of Antarctica have parts of the gut preserved by infilling matrix, showing that both mouth ad anus were located near the cone aperture. Muscle scars in other hyolith shells indicate that the animal had a series of dorsoventral and longitudinal, or longitudinal and circular muscles, which operated through a hydrostatic skeleton to protract and retract the head, to open and close the operculum, and to move the appendages. Although the shell form and skeletal ultra-structure of hyoliths are of a molluscan type, the muscle insertions suggest that the hyolith cone is not homologous with the dorsal exoskeleton of primitive molluscs. Hyoliths probably constitute a small extinct branch of phylum size, related to the Mollusca and the Sipunculoidea. All three groups may have had common ancestors in the late Precambrian.     

贵州剑河寒武系凯里组脊状单臂螺(Haplophrentis carinatus)埋藏特征及与始海百合共生关系探讨

贵州剑河寒武系凯里组软舌螺化石丰富、埋藏形式多样,其中以脊状单臂螺Haplophrentis carinatus化石数量最多、保存最为完整。过去有关学者对凯里组单臂螺化石的研究主要集中在化石分类学方面,对脊状单臂螺埋藏特征及与其他生物共生关系缺乏深入探讨。本文对324块脊状单臂螺化石标本进行系统研究对比后发现,凯里组脊状单臂螺口盖化石埋藏形式有四类:口盖单独保存、口盖以内模或外模化石形式保存、口盖与锥壳完全绞合保存、口盖与锥壳不完全绞合保存;附肢保存较少;锥壳多以内模化石形式保存;脊状单臂螺内模化石与印痕化石上普遍出现类似软体保存形成的三分叉结构,这类三分叉结构属于后期埋藏因素造成的次生结构。文中还对脊状单臂螺与始海百合共生关系进行探讨,并将二者共生关系归为偏害共生。     

Hyolith guts in the Cambrian of northern Australia -turning hyolithomorphs upside down

A single specimen of the early Middle Cambrian (Ordian) orthothecimorph hyolith Gudugu-wan hardmani with preserved gut filling has been discovered in the Montejinni Limestone, Wiso Basin, northern Australia. This represents only the eighth such occurrence worldwide. The specimen is unique among orthothecimorphs in retaining the gut filling in combination with the external mould of the operculum and invites revision of the accepted soft-part recon struction in this group, whereby the opercular lateral processes are now recognized to be ventral. Thus orthothecimorph conchs, if horizontal in life, would have been oriented ventral side down, while hyolithomorphs adopted the reverse orientation: horizontal, ventral side up.     

Shell composition of Terreneuvian tubular fossils from north‐east Sichuan,China

Identification of the primary constituents of small shelly fossil (SSF) shells is important for explaining the evolution of SSF faunas. The characteristics and constituents of Terreneuvian tubular SSFs found in north‐east Sichuan, China, are revealed using light microscopy, scanning electron microscopy and energy dispersive X‐ray spectroscopy. Petrographic thin sections indicate that the chemical composition of the shells is mainly calcium carbonate with smaller amounts of phosphorus, silica and pyrite. Most of the tubular shells composed of calcium carbonate have a distinct layered structure. Evidence of replacement of the original shell by phosphatization, pyritization and silicification, and recrystallization of calcium carbonate have been found, all of which destroyed the shell's original layered structure. Most fossils treated with acetic acid are phosphatic casts or steinkerns, with some preserving organic textures of the shell as phosphatic casts. We conclude that the Terreneuvian tubular SSFs from north‐east Sichuan were originally composed mainly of calcium carbonate; indeed, most Terreneuvian small skeletal fossils appear to have had an originally calcareous composition. The fossil casts or internal core fossils are composed of phosphate, which is related to local taphonomic processes.     

Aragonitic dendritic prismatic shell microstructure in Thracia (Bivalvia,Anomalodesmata)

The shells of most anomalodesmatan bivalves are composed of an outer aragonitic layer of either granular or columnar prismatic microstructure, and an inner layer of nacre. The Thraciidae is one of the few anomalodesmatan families whose members lack nacreous layers. In particular, shells of members of the genus Thracia are exceptional in their possession of a very distinctive but previously unreported microstructure, which we term herein “dendritic prisms.” Dendritic prisms consist of slender fibers of aragonite which radiate perpendicular to, and which stack along, the axis of the prism. Here we used scanning and transmission electron microscopical investigation of the periostracum, mantle, and shells of three species of Thracia to reconstruct the mode of shell calcification and to unravel the crystallography of the dendritic units. The periostracum is composed of an outer dark layer and an inner translucent layer. During the free periostracum phase the dark layer grows at the expense of the translucent layer, but at the position of the shell edge, the translucent layer mineralizes with the units typical of the dendritic prismatic layer. Within each unit, the c‐axis is oriented along the prismatic axis, whereas the a‐axis of aragonite runs parallel to the long axis of the fibers. The six‐rayed alignment of the latter implies that prisms are formed by {110} polycyclically twinned crystals. We conclude that, despite its distinctive appearance, the dendritic prismatic layer of the shell of Thracia spp. is homologous to the outer granular prismatic or prismatic layer of other anomalodesmatans, while the nacreous layer present in most anomalodesmatans has been suppressed.     

Shell structure of <Emphasis Type="Italic">Kutorgina</Emphasis> Billings (Brachiopoda,Kutorginida)

Shell structure in members of the genus Kutorgina, Lower Cambrian brachiopods with calcareous shells and primitive articulation, is described for the first time. This type of shell structure resembles that of the Lower Cambrian mollusks and hyoliths rather than of brachiopods.     

Hyolitha: status of the phylum

Hyoliths are operculate calcareous shells found in Palaeozoic rocks. Runnegar et al. (1975) suggested that they be referred to a new phylum (Hyolitha) but Marek & Yochelson (1976) and Dzik (1978) preferred to regard them as an extinct class of the Mollusca. Since the hyolith cone is not easily homologized with the monoplacophoran shell, the exoskeletons of the shelled Mollusca and the Hyolitha appear to have developed independently. Reconstructions of the anatomy of hyoliths indicate that it is unlikely that both groups shared a common molluscan ancestor. Therefore, hyoliths are probably not molluscs. Previous reconstructions of articulated hyolithids have suggested that left and right appendages (helens) curved dorsally. Crushed articulated specimens from the Burgess Shale indicate that this conclusion is incorrect; hyoltthid helens seem to have curved ventrally when the animals were alive.     

New data on molluscs and their shell microstructures from the Middle Cambrian Gowers Formation, Australia

Abstract: Numerous new cases of preserved shell microstructure were discovered in molluscs from the Middle Cambrian Gowers Formation (Ptychagnostus atavus/Peronopsis opimus Zone, Floran Stage) in the Georgina Basin, Australia. The new data provide further evidence that, by the Middle Cambrian, molluscan shell microstructures were diverse, and many molluscs had a complex shell with multiple types of shell microstructure. In addition, many new occurrences of laminar microstructures are described herein. For many, the nature of these laminar microstructures is not known, but in three species the microstructure is foliated calcite, and in at least two the microstructure is more likely to have been calcitic semi‐nacre, a type of microstructure known in brachiopods and bryozoans but unknown in modern molluscs. This commonality among these three closely related lophotrochozoans underscores a similar mechanism of biomineralization. Moreover, these observations suggest a prevalence of calcite‐shelled lineages among molluscs from the Middle Cambrian, a time of calcite seas. In addition, the broad occurrence of laminar, nacre‐like microstructures in many of these fossils reveals how widespread these strong (fracture‐resistant) microstructures were in Middle Cambrian molluscs. Additionally, a few specimens of Yochelcionella preserve imprints of a bilaterally symmetrical pair of muscle scars. New taxa described here include Corystos thorntoniensis gen. et sp. nov., Yochelcionella snorkorum sp. nov., Yochelcionella saginata sp. nov., and Anhuiconus? agrenon sp. nov.     

陕南地区下寒武统筇竹寺阶软舌螺化石

描述陕南地区下寒武统筇竹寺阶软舌螺化石６属８种,其中有５新属（Ningqiangethus gen.nov.,Bilgulitheca gen.nov.,Xixiangethes gen.nov.,Paranicrocornus gen.no.,Inflatatheca gen.nov.),新种,１相似种,１未定种。从组织结构和面貌上论述筇竹寺阶软舌螺和梅树村阶软舌螺匠区别,筇笔直寺阶的软舌螺不仅个体增大、背腹分异、口唇发育、壳体与口盖常共存,而且属种分异度大,为进一步研究软舌螺动物的早期演化和解决下寒武统生物地层划分为对比提供重要的实证材料。     

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.     

Wandel der Vorstellungen von der Frühevolution der Mollusken,besonders der Gastropoda und Cephalopoda

The ancestors of the molluscs are still unknown. Arguments in favour of flatworm-relationship are just as valid as those proposing segmented annelid-like worms as their closest relatives. The earliest molluscs lived before the onset of the Cambrian; but only at the end of this period the now existing classes of conchifers made their first appearance. Early and Middle Cambrian molluscan fossils are problematic. The hyoliths, for example, show relations to such fossils asSalterella andVolborthella, while closest living relatives can be found among tube-building annelids. Scars produced by tissue attached to the shell are of no use in the reconstruction of molluscan phylogeny, but a very useful tool in the analysis of the function regarding the body in interaction with its shell. The ontogenesis of the shell and muscles attached to it in recentFissurella (archaeogastropod) is presented as an example. Well known ontogenies in recent molluscs can aid to the reconstruction of fossils, demonstrated by some fossil gastropods. The interaction of soft and hard tissues, the function and structure of the shell of recent molluscs enables us to interpret the fossil forms, as for example the Lower Devonian coleoid cephalopods from the Hunsrück Schiefer. Reconstruction of the course of evolution is only possible, if information on living molluscs is integrated into historical data provided by the fossils.     

Shell structure, ontogeny and affinities of the Lower Cambrian bivalved problematic fossil Mickwitzia muralensis Walcott, 1913

Exceptionally preserved carbonate- and shale-hosted Mickwitzia muralensis from the Lower Cambrian Mural Formation, southern Canadian Rocky Mountains, complement one another to yield an unusually complete account of its ontogeny, ecology and phylogenetic relationships. The shell of M. muralensis is composed of dense phosphatic layers interspersed with porous organic-rich layers. At the insertion of shell-penetrating tubes, shell layers deflect inwards to produce inwardly pointing cones. The tubes are interpreted as having hosted setae that were secreted by outer-epithelial follicles. Follicular setae also occurred at the mantle margin, where they were oriented within the plane of the shell as in modern brachiopods. During ontogeny, the initial setae oriented in the plane of the shell occurred before the first shell-penetrative setae. In the juvenile and early-mature stages of shell secretion, a posterior opening was present between both valves and was used for the protrusion of an attachment structure. In the late-mature shell, this opening became fixed in the ventral valve. Based on the posterior margin and the shell microstructure, a close relationship between Mickwitzia and the paterinids is proposed with differences interpreted as heterochronic. The shell-penetrative setal apparatus of M. muralensis is distinct from that previously described of Micrina, though both types are conceivably homologous to adult and juvenile setae of modern brachiopods.     

Pyritized preservation of chancelloriids from the Cambrian Stage 3 of South China and implications for biomineralization

The enigmatic Cambrian animal chancelloriids were discovered in a wide range of taphonomic settings; however, preservation of biomineralized sclerite microstructure was solely known from secondarily phosphatized skeletal remains. Here, we investigate a uniquely pyritized chancelloriid from the lower Cambrian Guojiaba Formation in southern Shaanxi Province, China, using a combination of advanced analytic techniques. Results of the energy dispersive spectroscopy (EDS), X-Ray Fluorescence (XRF), and Raman spectrum show that the sclerites and scleritomes are preserved as pyritized internal moulds with a calcitic outer layer. The outer layer enveloping the internal moulds likely represents the recrystallized counterpart of the original biomineralized sclerite wall. Distinctive fibrous microstructures are discovered in the sclerites, which echo the features seen in the phosphatized fossils of chancelloriids. The typical microstructure, along with the recrystallized calcite, corroborate the interpretation that chancelloriid sclerites were originally constructed by fibrous aragonite. The stability of the microstructure and mineral composition in both carbonate and siliciclastic backgrounds indicate that chancelloriids were adapted to exploit aragonitic fibres to build their skeletons regardless of the change of their living environments.     

A first report of Sphenothallus Hall, 1847 in the Cambrian of Variscan Europe

Ten specimens of two phosphatic fossils have been recently discovered in lower and middle portions of Middle Cambrian Jince Formation in the Czech Republic. They are attributed to the genus Sphenothallus Hall, 1847 and described as two separate species; comparatively small conchs are described as S. kozaki sp. nov., the much larger specimens characterized by its smooth and partly flexible organo-phosphatic walls of shell are determined as ?S. kordulei sp. nov. Sphenothallus is known to range from Cambrian to Permian and accommodates numerous species. However, its Cambrian distribution is considerably restricted. Generally rare specimens have been described from Lower to Middle Cambrian of Laurentia and from the Lower Cambrian of Gondwana and peri-Gondwana. The new record of Sphenothallus from the Jince Biota represents a notable extension of their geographic range.     

Gracilitheca astronauta n. sp. and Nephrotheca sophia n. sp. (Hyolitha,Orthothecida) from the Cambrian of Czech Republic

Two orthothecid hyoliths, Gracilitheca astronauta n. sp. and Nephrotheca sophia n. sp. are described from the “middle” Cambrian Buchava Formation of the Skryje-Tý?ovice Basin in the Czech Republic. The new forms are based on about thirty well-preserved external and internal moulds of both conchs and opercula. Gracilitheca astronauta n. sp. is classified under the family Gracilithecidae Sysoev, 1972 and may be distinguished by clearly triangular cross-section of the conch, whereas Nephrotheca sophia n. sp. is characterized by transversally elongated cardinal processes reaching the lateral opercular margin and is combined with the family Orthothecidae Sysoev, 1958 emended Malinky, 2009. Several specimens of G. astronauta show operculum preserved inside of the conch. Stratigraphic ranges and geographic distributions within the Skryje-Tý?ovice Basin are established for both taxa.