Scleritome construction,biofacies, biostratigraphy and systematics of the tommotiid Eccentrotheca helenia sp. nov. from the Early Cambrian of South Australia

Abstract: Large collections of Eccentrotheca helenia sp. nov. from the lower Cambrian Wilkawillina and Ajax limestones in the Arrowie Basin, South Australia, contain abundant low, cap‐shaped and high, laterally compressed isolated sclerites in addition to partially articulated tubular specimens. The scleritome of Eccentrotheca helenia sp. nov. is fully described for the first time and shown to be formed by ontogenetic fusion of sclerites into successively stacked sclerite rings, forming a larger, tubular structure. The apical termination of the tube is highly variable, but is primarily constructed by low, cap‐shaped sclerites and characterised by a central aperture of variable inclination. The adapical portion of the tube is predominantly constructed by high, laterally compressed sclerites, but individual sclerite rings can contain both cap‐shaped and laterally compressed sclerites along with sclerites of intermediate morphology. The apical aperture presumably housed organic structures for attachment to a hard substrate, but the scleritome also occasionally preserves small lateral perforations between fused sclerites, which may have served to stabilise the scleritome by providing additional points of anchorage. In the Arrowie Basin, E. helenia is found in association with archaeocyath‐microbial‐spongiomorph‐dominated bioherms and most likely inhabited pendant or cryptic habitats within these bioherms. Eccentrotheca‐like sclerites form an integral part of the scleritomes of many tommotiids which may confuse taxonomic analysis. Sclerites previously assigned to ‘E.’guano, consistently occur together with sclerites of Kulparina rostrata in stratigraphic intervals consistently older than strata hosting E. helenia. Rare fused specimens indicate that the sclerites of K. rostrata and ‘E.’guano belong to the same scleritome.     

A Cambrian tommotiid preserving soft tissues reveals the metameric ancestry of lophophorates

1. Download : Download high-res image (342KB)
2. Download : Download full-size image

     

Redescription of Hexaconularia He and Yang, 1986 (Lower Cambrian, South China): implications for the affinities of conulariid-like small shelly fossils

Abstract:  Hexaconularia , a Lower Cambrian small shelly fossil (SSF) that has been allied with conulariids and scyphozoan cnidarians, is redescribed and refigured. A salient feature of this monospecific genus is the presence of distinct apical and abapical regions. The apical region probably represents an embryonic shell that apparently lacked a basal attachment structure. Comparisons of this feature with the apical end of the smallest known conulariids and with conulariids terminating in an apical wall (schott) reveal substantial differences in structure and ornamentation. Differences in apical anatomy between conulariids and Arthrochites , possibly the nearest SSF relative of Hexaconularia , are also apparent. Comparisons of Hexaconularia with Punctatus , an SSF taxon showing distinct apical and abapical regions in both posthatching specimens and prehatching embryos, suggest that the early development of Hexaconularia was direct. These results have important implications for hypotheses of a conulariid/scyphozoan affinity for Hexaconularia and its possible SSF relatives, and they suggest that Hexaconularia -bearing strata may yield prehatching embryos of this genus.     

Beyond the Burgess Shale: Cambrian microfossils track the rise and fall of hallucigeniid lobopodians

Burgess Shale-type deposits are renowned for their exquisite preservation of soft-bodied organisms, representing a range of animal body plans that evolved during the Cambrian ‘explosion’. However, the rarity of these fossil deposits makes it difficult to reconstruct the broader-scale distributions of their constituent organisms. By contrast, microscopic skeletal elements represent an extensive chronicle of early animal evolution—but are difficult to interpret in the absence of corresponding whole-body fossils. Here, we provide new observations on the dorsal spines of the Cambrian lobopodian (panarthropod) worm Hallucigenia sparsa from the Burgess Shale (Cambrian Series 3, Stage 5). These exhibit a distinctive scaly microstructure and layered (cone-in-cone) construction that together identify a hitherto enigmatic suite of carbonaceous and phosphatic Cambrian microfossils—including material attributed to Mongolitubulus, Rushtonites and Rhombocorniculum—as spines of Hallucigenia-type lobopodians. Hallucigeniids are thus revealed as an important and widespread component of disparate Cambrian communities from late in the Terreneuvian (Cambrian Stage 2) through the ‘middle’ Cambrian (Series 3); their apparent decline in the latest Cambrian may be partly taphonomic. The cone-in-cone construction of hallucigeniid sclerites is shared with the sclerotized cuticular structures (jaws and claws) in modern onychophorans. More generally, our results emphasize the reciprocal importance and complementary roles of Burgess Shale-type fossils and isolated microfossils in documenting early animal evolution.     

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.     

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.     

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.     

The Cambrian brachiopod fauna from the first-trilobite age Shuijingtuo Formation in the Three Gorges area of China

The Yangtze platform of South China offers evidence within its Ediacaran–Cambrian geological record of the Cambrian explosion and diversification events in metazoan history. To understand the explosive radiation of animals and the environments in which it took place, the basal Cambrian fauna succession of the Aijiahe section in the Three Gorges area, western Hubei Province, has been studied, revealing the earliest brachiopod fauna (Tsunyidiscus trilobite Zone) in this region, which was dominated numerically by acrotretoids. This is accompanied by abundant skeletal fossils including minute well-preserved phosphatized archaeocyath cups and an assortment of abundant sponge spicules, chancelloriids, mollusks, hyoliths, and bradoriids, retrieved by acid-etching limestone interbeds in the black shale-dominated Shuijingtuo Formation (Series 2). The brachiopods comprise two species of acrotretoids, two types of botsfordiids (Botsfordiidae gen. et sp. indet. A and B), and four species of linguloids. Of the latter, Spinobolus popovi n. gen. n. sp. is strikingly distinctive and typified by spine-like ornamentation seen for the first time in the lower Cambrian; the remaining three linguloid genera, Palaeobolus, Eoobolus, and Lingulellotreta, have a trans-paleocontinental distribution. The Three Gorges Shuijingtuo brachiopod assemblage differs from that of the upper Atdabanian Stage (Cambrian Stage 3) in Siberia and South China, but shows great similarities with those discovered in the Tsanglangpuan (equivalent to Botoman or Stage 4) Stage of eastern Yunnan Province, Siberia, and South Australia, suggesting a much more prolonged sedimentary hiatus in basalmost Shuijingtuo Formation of the Three Gorges area than previously expected. The presence of such unconformities provides a caveat to stable isotope-based correlations that involve a number of discussions of global ocean geochemical changes across the time interval that witnessed Cambrian explosion of metazoans.     

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.     

BASAL CAMBRIAN MICROFOSSILS FROM THE YURTUS AND XISHANBLAQ FORMATIONS (TARIM, NORTH-WEST CHINA): SYSTEMATIC REVISION AND BIOSTRATIGRAPHIC CORRELATION OF MICRHYSTRIDIUM-LIKE ACRITARCHS

Abstract:  Micrhystridium -like acritarchs are widely distributed in basal Cambrian cherts and phosphorites in South China. This paper describes similar acritarchs from the basal Cambrian Yurtus and Xishanblaq formations in Tarim, north-west China. The taxonomy of these acritarchs is revised. The basal Cambrian acritarch assemblage in Tarim and South China is characterized by three genera: Asteridium Moczydłowska, Heliosphaeridium Moczydłowska and Comasphaeridium Staplin, Jansonius and Pocock. This assemblage is named the Asteridium - Heliosphaeridium - Comasphaeridium (AHC) acritarch assemblage. In both South China and Tarim, the AHC acritarch assemblage is associated with the tubular microfossil Megathrix longus Yin L. and the small shelly fossil Kaiyangites novilis Qian and Yin G. This assemblage also occurs in the Lower Tal Formation in the Lesser Himalaya. Correlation with small shelly fossil (SSF) assemblages indicates that the AHC assemblage is restricted to the Meishucunian Stage, and possibly to the lower Meishucunian ( Anabarites trisulcatus - Protohertzina anabarica and Siphogonuchites triangularis - Paragloborilus subglobosus SSF assemblages). The AHC assemblage is broadly similar to the Asteridium tornatum - Comasphaeridium velvetum (acritarch) Zone in the East European Platform, which is considered to be Nemakit-Daldynian (and possibly Tommotian) in age.     

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.     

Lapworthellids and other skeletonised microfossils from the Cambrian Stage 3 of the northern Montagne Noire,southern France

The Montagne Noire (southern France) possesses one of the most complete Cambrian successions in the western peri-Gondwana margin and might provide a good stratigraphic reference for both regional charts and international correlations. However, to date, the lower Cambrian succession of the northern Montagne Noire has been supposed to be devoid of biostratigraphically significant fossils. The complex tectonostratigraphic framework of the area (a range divided into Axial Zone, northern and southern Montagne Noire) exacerbated problems related to regional correlations and palaeogeographic reconstructions. As a result, the chronostratigraphic context of the lower Cambrian of northern Montagne Noire is still uncertain and stratigraphic reports have broadly relied on putative lithostratigraphic correlations with the southern Montagne Noire. The purpose of this study is to characterise, for the first time, the fossil record of carbonate beds and lenses of the northern Montagne Noire occurring at the top of the siliciclastic-dominated Marcory Formation, in order to provide regional bio- and chronostratigraphic constraints on lower Cambrian strata. Moreover, this study is aimed at improving international chronostratigraphic correlation. Carbonate beds and lenses cropping out along the Orque Cliff, in the Mélagues thrust slice, were investigated. They yielded a faunal assemblage constituted of molluscs (Igorella cf. ungulata and Igorella moncereti n. sp.), hyoliths (Conotheca brevica), chancelloriids (Archiasterella cf. pentactina and Allonnia sp.) and tommotiids (Lapworthella rete). L. rete is recorded in upper Meishucunian (Cambrian Stage 3) strata of the Yangtze Platform (South China) where it is used as index fossil of the Cambrian Stage 3 Sinosachites flabelliformis–Tannuolina zhangwentangi Assemblage Zone. Therefore, the presence of this tommotiid provides evidence that the studied carbonate beds and lenses are Cambrian Age 3 (Atdabanian according to the Siberian chart). The upper part of the Marcory Formation in the Mélagues slice, dated as Cambrian Stage 3, might represent a lateral equivalent of the mixed (carbonate-siliciclastic) Pardailhan Formation defined in the southern Montagne Noire.     

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.     

RECONSIDERATION OF THE PHYLOGENETIC AFFINITIES OF ZHIJINITIDS

Isolated sclerites are an important component of the Early Cambrian small shelly fos-sils(SSF) ,accounting for one-third of both the total number of taxa and the total numberof specimens.Estimating the phylogenetic affinities of these sclerites is one of the mostdifficult problems in the analysis of the SSF.Sclerites referred to such taxa as the halkieri-ids,siphogonuchitids,chancelloriids,tommotiids,zhijinitids and cambroclavids generallyhave been interpreted as Problematica.As recently a…     

Ontogeny of the eodiscoid trilobite Tsunyidiscus acutus from the Lower Cambrian of South China

Abstract: The morphology and ontogeny of the eodiscid trilobite Tsunyidiscus acutus Sun is described on the basis of numerous calcified specimens collected from the Lower Cambrian Shuijingtuo Formation in Yichang and Changyang, Hubei Province, South China. An ontogenetic series is established based on the articulated material including the previously unknown protaspides and meraspides (degrees 0 and 1). The material, revealing some prominent morphological changes such as the number of pygidial axial rings, genal spines retained throughout ontogeny and pygidial pleurae from furrowed to unfurrowed, enables a discussion on the trunk segmentation schedule, indicating that somitogenesis and tagmosis occurred independently during the ontogenetic development of T. acutus.     

再论织金钉类(Zhijinnids)的生物亲缘

寒武纪最早期层位中获得的离散骨片化石Halkieria和Microdictyon,已在较高层位的特异埋藏化石群中找到了它们的完整生物躯体,它为研究离散骨片化石的自然分类和生物亲缘提供了直接证据。本文通过寒武纪最早期钉状化石的形态、对称性、壳质成分、矿化特征的对比,提出了离散骨片化石织金钉(Zhijinites)和拟织金钉(Parazhijinites)与澄江动物群中的怪诞虫(Hallucigcnia)和爪网虫(Onychodictyon)等体表上的钉状物有近亲关系,它们可能同属于多足缓步类(Tardipolypoda)。     

陕西镇巴下寒武统水井沱组小壳化石新属种

本文描述了陕西镇巴下寒武统水井沱组中的小壳化石３新属５新种。样品采自镇巴小洋坝曾家坡,共生化石有三叶虫、古介形虫、海绵骨针和大量软舌螺等。化石组合面貌表明其地质时代属筇竹寺期。另外,在峡东水井沱组中发现了微网虫（Ｍｉｃｒｏｄｉｃｔｙｏｎｓｐ．）化石骨片（图版Ｉ,图１８,１９）,可与云南筇竹寺组及陕南水井沱组对比,说明Ｍｉｃｒｏｄｉｃｔｙｏｎ广泛分布于扬子地台区。     

陕西洛南下寒武统辛集组疑难管状化石桶螺(Cupitheca)

本文对采自陕西洛南上湾剖面下寒武统辛集组疑难管状化石Cupitheca进行了研究,系统厘定和描述了两个种,分别为Cupitheca holocyclata和Cupitheca costellata,并对Cupithecidae科进行了重新修订。本文所描述标本的壳体多为次生磷酸盐化保存,壳体表面纹饰保存较好;C.holocyclata管体表面为横向纹饰,C.costellata管体表面为纵向纹饰。C.holocyclata全球广布,如南澳大利亚、南极乔治王岛、格陵兰东北部、加拿大纽芬兰等地皆有报道,产出于下寒武统中上部地层,具有一定的洲际地层对比意义。本文为C.holocyclata在中国首次发现。C.costellata仅发现于华北,即本文所研究的洛南辛集组及其同期地层安徽淮南与霍邱的雨台山组。     

陕西南部早寒武世原始锥石类carinachitids两新种记述

对陕西宁强宽川铺地区灯影组宽川铺段原始锥石类carinachitids两个新种Emeiconularia amplicanalis sp.nov.和Carinachites platus sp.nov.进行了记述。外表形态多样、个体大小悬殊的两辐对称、三辐对称和四辐对称的carinachitids系列标本的发现,揭示了陕南地区早寒武世丰富多样的carinachitids生命世界。三辐射对称的Emeiconularia在四川峨眉及陕西南部的震旦-寒武系界线附近都有分布,因而对前寒武纪—寒武纪跨界地层具有划分对比意义。两新种的发现无疑增加了我们对早期锥石类多样性的认识,为证实多样性的刺细胞动物在早寒武世早期已经存在的假说提供了新线索。