SKELETAL MICROSTRUCTURE INDICATES CHANCELLORIIDS AND HALKIERIIDS ARE CLOSELY RELATED

Abstract: Chancelloriids are problematic, sac‐like animals whose sclerites are common in Cambrian fossil assemblages. They look like sponges, but were united with the slug‐like halkieriids in the group Coeloscleritophora Bengtson and Missarzhevsky, 1981 based on a unique mode of sclerite construction. Because their body plans are so different, this proposal has never been well accepted, but detailed study of their sclerite microstructure presented here provides additional support for this grouping. Both taxa possess walls composed of a thin, probably organic, sheet overlying a single layer of aragonite fibres orientated parallel to the long axis of the sclerite. In all halkieriids and in the chancelloriid genus Archiasterella Sdzuy, 1969 , bundles of these fibres form inclined projections on the upper surface of the sclerite giving it a scaly appearance. On the lower surface of the sclerite, the projections are absent. This microstructure appears to be unique to chancelloriids, halkieriids, and their relatives, siphogonuchitids and sachitids. (The sclerites of another putative halkieriid relative, Wiwaxia Walcott, 1911 , are unmineralized, making direct comparisons impossible.) Thus, similarity both at the level of sclerite construction and the level of sclerite microstructure suggests that chancelloriid, halkieriid, sachitid, and siphogonuchitid sclerites are homologous. The difference in chancelloriid and halkieriid body plans can be resolved in two ways. Chancelloriids either represent a derived condition exhibiting complete loss of bilaterian characters or they represent the ancestral condition from which the bilaterally symmetric halkieriids, and the Bilateria as a whole, derived. The latter interpretation, proposed by Bengtson (2005) , implies that coeloscleritophoran sclerites (‘coelosclerites’) are a plesiomorphy of the Bilateria, lost or transformed in descendent lineages. Given that mineralized coelosclerites appear in the fossil record no earlier than c. 542 Ma, this in turn implies either that the Ediacaran record of bilaterians has been misinterpreted or that coelosclerite preservability increased at the beginning of the Cambrian Period. The former is difficult to reconcile with Ediacaran trace and body fossil evidence, but the latter may be possible, reflecting either independent mineralization of organic‐walled sclerites in chancelloriids and halkieriids or the opening of a taphonomic window that favours coelosclerite preservation.     

Chancelloriid sclerites from the Lower Cambrian (Meishucunian) of eastern Yunnan,China, and the early history of the group

Chancelloriids are a group of enigmatic sessile animals that are covered with sclerites shaped as rosettes of spines, producing an appearance like that of a barrel cactus. They are known only from Cambrian rocks. Isolated sclerites of chancelloriids are widespread in small shelly faunas, but they have proven difficult to treat taxonomically due to the variation within and between individual animals. We report on large samples of chancelloriid sclerites from the Meishucunian (pre‐trilobitic Lower Cambrian) of eastern Yunnan Province, China, including material from the Dahai Member of the Zhujiaqing Formation (the Zhujiaqing section, Dahai, Huize County) and the Shiyantou Formation (the Xiaotan section, Yongshan County). The material from the Dahai Member appears to come from a single species, Chancelloriella irregularis. Statistical analysis of morphotype co‐occurrences in different samples suggests that several species are represented in the material from the Shiyantou Formation, which we herein tentatively place in four species, Allonnia erromenosa, Allonnia tetrathallis, Archiasterella charma sp. nov. and Archiasterella cf. pentactina. We suggest therefore that careful study of large collections of isolated chancelloriid sclerites permits the identification of different species, opening the possibility of their use in evolutionary or stratigraphical studies. In particular, in South China, it seems that Chancelloriella irregularis is characteristic of rocks of the Paragloborilus subglobosus – Purella squamulosa Assemblage Zone and the Watsonella crosbyi Assemblage Zone, whilst Archiasterella and Allonnia first appear in the overlying Sinosachites flabelliformis – Tannuolina zhangwentangi Assemblage Zone. C. irregularis is thus amongst the oldest well‐characterized chancelloriids, and the irregularity and poor organization of its sclerites relative to those of younger forms support the hypothesis that chancelloriid sclerites are compound structures arising from the fusion of originally separate elements, perhaps homologous with sclerites of halkieriids and other coeloscleritophorans. The similarity between the arrangement of rays in a chancelloriid sclerite and the arrangement of bubbles in small bubble clusters suggests that in many cases, aspects of the form of chancelloriid sclerites are the result of simple physical processes.     

寒武纪叶足动物链状心网虫躯干骨板功能形态学研究*

链状心网虫(Cardiodictyon catenulum)是寒武纪澄江生物群中较为常见的叶足动物, 其躯干骨板的形态一直存在较大争议。本项研究显示该叶足动物的躯干具有一纵列马鞍状骨板, 每个体节具有单一的骨板, 每个骨板从背部延伸至躯干两侧。因此该叶足动物的骨板与其余具有成对或环状分布的躯干骨板的叶足动物存在显著的区别。骨板中央存在一个圆锥形凸起, 骨板边缘显著增厚且呈锯齿状。链状心网虫的躯干骨板边缘增厚和鞍点的凸起提升了整体的刚性; 锯齿状边缘也指示了更优的保护及防御功能。本项研究还发现链状心网虫骨板表面具有显微筛状结构, 筛孔被充填并保存为众多等距离分布的黄铁矿化瘤突, 与微网虫、怪诞虫及爪网虫躯干骨板的显微网状结构类似, 可能为纤细乳突的出口。本文认为: 寒武纪叶足动物的躯干骨板普遍具有显微筛状结构; 这些骨板除了保护与防御, 可能还有感觉的功能。每个躯干体节对应骨板为寒武纪叶足动物的独有衍征, 可能为背骨板的早期形态, 也可能由成对骨板愈合而来。     

Chancelloriid sclerites from the lowermost Cambrian of North China and discussion of sclerite taxonomy

Chancelloriid sclerites from the lowermost Cambrian Xinji Formation (Series 2, Stage 3), southern margin of the North China platform, are systematically described. Thousands of isolated sclerites from three sections are assigned to three genera and four species, including Chancelloria cf. eros, Allonnia tripodophora, Archiasterella pentactina, and Ar. tetraspina. To accurately document the taxonomic significance of the sclerite structure, modified formulas (m+nC, m+nA, and m+0) are put forward to represent the full series and variation of sclerite forms. Based on the sclerite construction, statistical analysis on the proportions of different sclerite forms in the rock samples and the composition of sclerites in previously described chancelloriid scleritomes, a new scheme for identification and classification of isolated sclerites reconciled within the framework of the complete scleritome, is proposed.     

A lower Cambrian protoconodont apparatus from the Placentian of southeastern Newfoundland

A cluster of three pyritized, naturally arranged, conoidal sclerites has been found on a bedding surface in the lower Cambrian Chapel Island Formation in southeastern Newfoundland. The elements are about 4 mm in length and are morphologically similar to protoconodonts but neither their original chemical composition nor internal structure is preserved. Arrangement of the elements suggests that they belonged to an apparatus that performed a grasping function. The specimen is described here as ProtohertzinaEfficacynadia n.sp. This is the oldest hitherto described protoconodont apparatus, apart from clusters of Mongolodus described from the lower Cambrian of the Lesser Himalaya. This paper adds to the scant knowledge of early protoconodonts that we consider to be the earliest biomineralized predators.     

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.     

Shell microstructures of the Early Cambrian Anabarella and Watsonella as new evidence on the origin of the Rostroconchia

Internal moulds of the laterally compressed monoplacophoran Anabarella plana Vostokova, 1962, and likely earliest rostroconch Watsonella Grabau, 1900, from the Early Cambrian of the Siberian Platform, show similar microstructures. The moulds are covered with a thin phosphatic crust replicating the inner morphology and microstructure of the shells. The shells were completely removed during etching of the samples in 10% acetic acid, except for some moulds of Watsonella sp., which retained an incompletely preserved and recrystallized wall. Three types of microtexture were found in moulds of Anabarella and Watsonella : polygonal, spiny and step-wise. The polygonal texture is well exposed in the apical area and dorsal margin and is interpreted to represent a prismatic outer layer. The polygons can be partially overlapped by spiny and stepwise textures that may represent a crossed-lamellar inner layer. Prisms and lamellae were first-order structural units, probably consisting of fibers. The similar shell microstructures of Anabarella and Watsonella , especially at the dorsal margins, support the hypothesis that a laterally compressed monoplacophoran such as Anabarella plana was the first evolutionary step from monoplacophorans towards Early Cambrian bivalves via the earliest rostroconch-like Watsonella.     

A comparative study of Lower Cambrian Halkieria and Middle Cambrian Wiwaxia

Two Cambrian lepidote metazoans known from different respective types of preservation have been compared in order to elucidate their biology and affinities. The widely distributed Lower Cambrian Halkieria is represented by isolated hollow sclerites, probably of originally calcareous composition. The Middle Cambrian Wiwaxia is known from the Burgess Shale as isolated sclerites (scales and spines) and as more or less complete individuals. Although Halkieria sclerites were mineralized and those of Wiwaxia were probably not, there are fundamental structural and morphological similarities between the two. Both bad an imbricating scaly and spiny armour consisting of hollow sclerites with a longitudinally fibrous structure. The sclerites did not grow, but were probably moulted during the course of ontogenetic growth. Halkieria and Wiwaxia are regarded as closely related. Both are referred to the Order Sachitida He 1980. The sclerite armour of Halkieria is reconstructed on the template provided by Wiwaxia. The interpretation of sachitid sclerites as protective armour is an alternative to the interpretation by Jell (1981, Alcheringa 5 )that sachitid sclerites were respiratory organs in an animal of probable annelid affinities. Sachitids are interpreted as sluggish, benthic deposit feeders that do not belong to any recognized phylum.     

Early Cambrian lobopodian sclerites and associated fossils from Kazakhstan

A low diversity fossil assemblage dominated by Mongolitubulus spines, from the Early Cambrian Geres Member of the Shabakhty Formation at Koksu, Malyi Karatau, Kazakhstan, enables identification of its sclerite variability within the scleritome. No other sclerotised structures could be matched with the spines, indicating that these were the only sclerotised part of the body and were periodically shed. The prominent spines of Mongolitubulus have a scale–like external ornamentation and fibrous internal structure. Possibly they armed the body of a lobopodian similar to Xenusion . The Mongolitubulus spines belong to a series of Cambrian phosphatic fossils, the other end members of which are sclerites of Microdictyon , the second most abundant component of the Koksu assemblage. The latter superficially resemble schizochroal trilobite eyes, but in life covered the dorso–lateral sides of the body segments, as is evident from complete specimens from the Chinese Chengjiang locality. A pair of sclerites of Microdictyon , representing uncompleted exuviation, from Koksu show that each hole originally contained a non–phosphatised lenticular body. This makes a compound eye nature of the Microdictyon sclerites likely, and even their homology with arthropod eyes cannot be excluded.     

A new mongolian species of the lower cambrian genus camenella and the problems of scleritome-based taxonomy of the tommotiidae

The tommotiidCamenella parilobata n. sp. is described on the basis of a collection of mitral and sellate sclerites from the Lower Cambrian Khairkhan »Formation« of western Mongolia. The detailed documentation of co-occurrence of sclerite types is essential for the taxonomic analysis of the problematic Tommotiidae and related groups; most »tommotiid« (sensu lato) taxa are still known only from a few isolated sclerites representing an unknown spectrum of variability. Until a detailed analysis of the scleritome composition and sclerite microstructure in representatives of all genera has been made, any suprageneric classification can only be speculative.     

湖北秭归茅坪镇下茶庄剖面石牌动物群初探*

报道湖北秭归长江以南茅坪镇下茶庄剖面发现的寒武系下部(第2统第4阶)石牌动物群。该剖面石牌组出露良好,顶底界线清楚,总厚度208m。初步采集结果表明该剖面具有较大的开采潜力,化石数量上以腕足动物和软舌螺为主,同时还包括数量较多的三叶虫、双瓣壳节肢动物、开腔骨类和少量的鳃曳动物等,其中的腕足动物保存良好,数量丰度较大,壳体多以密集立体保存的内模形式出现,这是石牌组腕足动物大量壳体化石的首次报道。化石采集发现鳃曳动物与腕足动物可在相同的层面同时出现。与澄江动物群鳃曳动物化石相比,下茶庄剖面鳃曳动物的环纹保存的立体程度明显较强,但还未见到任何吻部和肠道结构保存。石牌动物群最早发现于长江以北宜昌三峡人家旅游风景名胜区内的王家坪剖面,但该地区石牌组覆盖严重,顶底未见出露,使王家坪剖面石牌动物群的后续研究存在较大困难。因此,石牌动物群在下茶庄剖面的发现和进一步采集和研究对深入揭示石牌动物群的组成、富集层位、群落结构和地理分布范围有重要的意义。     

我国西南地区早寒武世小壳化石Sinosachites

采自四川峨嵋和云南会泽的磷酸盐化的Sinosachites骨片显示出中央沟和侧沟结构,并区别出三种不同类型的骨片(palmate,cultrate and siculate)。这一发现证实,产于澳大利亚的Thambetolepis(Jell,1981)是Sinosachites的晚出同义名。基于上述结构,该产地一些原定为Sachites和Halkieria的标本亦应归入Sinosachites。在Sinosachites属中确认出两个种,即产于华南的Sinosachites flabelliformis He和产于澳大利亚的Sinosachites delicatus(Jell)。在前一种中,左右型对称骨片的存在证实该动物两侧对称。骨片的基部通常保存为磷质内核,表明它比骨片壁更容易磷酸盐化,在组织学上它与骨片壁不同。骨片的基部可能原为有机质或部分矿化。在Sinosachites中,无帽状和束状骨片可能表明在coeloscleritophorans中halkieriids是一个介于wiwaxids和siphogonuchitids之间的类群。     

Plywood‐like shell microstructures in hyoliths from the middle Cambrian (Drumian) Gowers Formation,Georgina Basin,Australia

Hyoliths are a group of Palaeozoic fossils with calcareous shells whose affinities remain controversial. As their shells were originally aragonitic, their fossils are usually coarsely recrystallized, and few data on their microstructure are available. We report hyoliths from the middle Cambrian (Drumian, Floran) Gowers Formation of the eastern Georgina Basin, Queensland. These are preserved as phosphatic internal moulds, often with the inner layers of the shell also partly replaced by phosphate. Microstructural details preserved by this early diagenetic phosphatization show that these hyolith conchs were originally composed of fibrous crystallites, c. 0.5 μm wide, parallel to one another and to the inner surface of the shell. In several species, the fibres are arranged in a plywood‐like structure composed of multiple lamellae with a different fibre orientation in each lamella: often they are transversely oriented (relative to the long axis of the conch) in the inner part of the wall and longitudinally oriented in the outer part. Opercula also show a microstructure of parallel fibres. The lamello‐fibrillar microstructure we report from hyoliths is reminiscent of microstructures of many Cambrian molluscs; that this microstructure is found in both conchs and opercula suggests that these structures are serial homologues of one another, and in this respect they resemble brachiopod valves. As with many other biological plywoods, the hyolith shell probably records self‐organization in a liquid‐crystal‐like organic matrix. This provided a straightforward way to construct a material that could resist stresses from different directions, offering an effective defence against predators.     

Formation and morphology of epidermal sclerites from a deep-sea hydrothermal vent solenogaster (Helicoradomenia sp., Solenogastres, Mollusca)

The deep-sea hydrothermal vent solenogaster Helicoradomenia is covered with calcium carbonate sclerites. Light and electron microscopy reveal varying morphologies of these sclerites. Many sclerites have hollow tips and/or are pitted and etched. Bacteria are found on and in sclerites. Initial sclerite formation occurs in an extracellular crystalline chamber formed by the invagination of a cuboidal basal cell of the columnar microvillus mantle epithelium. As the sclerite grows, it fills the crystalline chamber resulting in direct contact with the microvilli of both the basal cell and neighboring secondary sclerite-forming cells. These cells shape a collar around the base of the growing sclerite. As growth continues, the sclerite-forming cells stretch around the sclerite forming a sheath in which the base of the sclerite resides. Mature sclerites grow through the cuticle into the external environment. The erosion pattern of sclerites reveals a less stable inner medullary region and a harder outer cortical region. This points to a secondary character state, where foremost hollow acicular sclerites develop into solid sclerites. This is in agreement with the systematic position of the genus Helicoradomenia within Simrothiellidae, a taxon typically with hollow sclerites.     

The tommotiid Kelanella and associated fauna from the early Cambrian of southern Montagne Noire (France): implications for camenellan phylogeny

A carbonate bed of the Pardailhan Formation, early Cambrian, southern Montagne Noire (southern France), provided microfossils such as Hyolithellus sp., Torellella cf. mutila and Torellella sp. along with numerous disarticulated pieces of composite skeletons such as valves of the brachiopod Eoobolus priscus and of the bradoriid Monceretia erisylvia, and chancelloriid sclerites (Chancelloria sp.). The assemblage also furnished a rich set of sclerites from the tommotiid Kelanella altaica. Five morphological variations of the latter have been identified. The presence of concentric ribs formed by distal inflation of selected shell laminae in Kelanella supports its assignment to the camenellans. More particularly, the presence of transverse structures within the internal cavity (septa) of Kelanella suggests a close relationship with the Lapworthellidae. However, the latter differ from Kelanella by the continuous morphological variation along their scleritome which is also composed of simple conical elements with uniform ornamentation. Several forms of Kelanella are similar to mitral and sellate sclerites of Camenella, whereas some other forms are comparable to Kennardia. The new material suggests that Kelanella occupies a transitional position between Lapworthellidae and the grouping of Tommotiidae and Kennardiidae. Such a phylogenetic position also implies that the number of sclerite morphotypes tends to decrease within the camenellan scleritome during evolution.     

Paterimitra pyramidalis from South Australia: scleritome,shell structure and evolution of a lower Cambrian stem group brachiopod

The tommotiid Paterimitra pyramidalis Laurie, 1986, is redescribed based on well‐preserved material from the lower Cambrian Wilkawillina, Wirrapowie and Ajax limestones of the Flinders Ranges, South Australia. The material shows that the scleritome of Paterimitra pyramidalis includes three sclerite morphotypes (S1, S2 and L). Detailed shell microstructure studies show striking similarities with both the paterinid brachiopod Askepasma toddense and the tommotiid Eccentrotheca helenia, which strengthens the suggested evolutionary link between tommotiids and brachiopods. Based on the partly articulated specimens and similarities in shell microstructure and sclerite morphology with Eccentrotheca, Paterimitra pyramidalis is reconstructed as a tube‐dwelling, epifaunal, sessile, filter‐feeder with an organic pedicle‐like attachment structure. The proposed reconstruction of the scleritome comprises a basal unit composed of one S1 and one S2 sclerite, as well as an unresolved number of L sclerites lining a coniform tubular structure.     

Middle Cambrian chancelloriids from the Argentine Precordillera

We report, for the first time, and systematically describe chancelloriid from the region, including Chancelloria bella, Allonnia cf. tintinopsis, Al. cf. erromenosa, Al. tripodophora, Archiasterella cf. coriacea, Ar. cf. hirundo, Ar. cf. tetraspina, and Ar. cf. fletchergrully, along with previously reported chancelloriid species (Chancelloria cruceana, C. eros, and Archiasterella sp.). The current chancelloriid fauna is similar in taxonomic composition to the Cambrian Burgess Shale-type biotas and to those of China and South Australia. Based on the sclerite construction, we found that the different articulatory facet types may be used to distinguish Allonnia from Archiasterella, and the number of rays is closely related with the arrangement of sclerite rays. The chancelloriid fauna is found stratigraphically between the Glossopleura trilobite zone of the restricted-shelf facies and the Oryctocephalus trilobite zone of the open-shelf facies, spanning the Wuliuan Stage (Delmaran/Topazan) of the Miaolingian Series, middle Cambrian. This study is significant for better understanding the diversity of this enigmatic and cosmopolitan group in the middle Cambrian warm platform of the Precordillera.     

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.     

云南永善肖滩剖面早寒武世Lapworthella (托莫特壳类)骨片化石

描述了采自云南永善肖滩剖面下寒武统石岩头组的托莫特壳类(tommotiids)骨片化石Lnpworthella rete Yue,1987,其骨片锥状或塔状,横向环脊明显,环脊尖棱状;骨片外表面具多边形网状纹饰,但无齿、瘤或纵脊等装饰;骨片可分为两种类型,即弯曲型和近直立型,前者数量明显多于后者。简要地讨论了托莫特壳类化石的分类及生物亲缘关系,认为目前为止,其仍为亲缘关系不明的化石类别。尚难断定所有的托莫特壳类骨片化石皆属于同一生物门类,因为难以证明将它们归人同类生物所依据的骨片性状是共同衍征(synapomorphy)还是由于趋同演化(convergent evolution)的结果;在缺乏躯干化石记录的情况下,离散骨片的生物亲缘关系研究较为困难。     

湖北房县下寒武统西蒿坪段钉形骨片化石

本文对采自湖北房县三座庵剖面下寒武统西蒿坪段的钉形骨片(nail-shaped)化石进行了研究,系统厘定和描述了2属2种,分别为Parazhijinites guizhouensis和Cambroclavus angxianensis;其中以C.fangxianensis骨片化石最为丰富,由三种类型的骨片组成,它们可能...