Extraordinary fossils reveal the nature of Cambrian life: a commentary on Whittington (1975) ‘The enigmatic animal Opabinia regalis,Middle Cambrian,Burgess Shale,British Columbia’

Harry Whittington''s 1975 monograph on Opabinia was the first to highlight how some of the Burgess Shale animals differ markedly from those that populate today''s oceans. Categorized by Stephen J. Gould as a ‘weird wonder’ (Wonderful life, 1989) Opabinia, together with other unusual Burgess Shale fossils, stimulated ongoing debates about the early evolution of the major animal groups and the nature of the Cambrian explosion. The subsequent discovery of a number of other exceptionally preserved fossil faunas of Cambrian and early Ordovician age has significantly augmented the information available on this critical interval in the history of life. Although Opabinia initially defied assignment to any group of modern animals, it is now interpreted as lying below anomalocaridids on the stem leading to the living arthropods. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

峡东地区寒武纪水井沱组下段双瓣壳类节肢动物系统古生物与个体发育研究

双瓣壳类节肢动物在全球寒武纪海洋中广泛分布,是寒武纪时期布尔吉斯页岩型化石生物群的重要组成类群和典型代表。它们的起源、演化及生态学研究是对早期后生动物演化研究的重要内容。本文详细描述了峡东地区寒武系第二统第三阶水井沱组下段两种双瓣壳类节肢动物,Caudicaella bispinata (Cui and Huo,1990) comb.nov.和Sunella grandis Huo,1965。依据新建的Caudicaella属征,对孙氏虫科的鉴定特征进行了修订。首次研究了孙氏虫科两属种壳瓣的生长发育模式,揭示了壳瓣高长比的等速生长和C.bispinata基刺相对于壳长的异速生长关系。元素扫描分析研究表明,长阳地区天柱山村剖面化石为碳质压膜保存。功能形态学研究认为,C.bispinata是远洋浮游型生物,其广泛的古地理分布有望为寒武纪早期全球生物地层对比提供新的化石依据。     

Oldhamia radiata: the trace of an undermat miner from the Cambrian Series 2 Xinji Formation,western Henan,North China

Oldhamia trace fossils represent a complex horizontal burrow system usually associated with microbial mats, occurring in the upper part of the Cambrian Stage 2 of the Terreneuvian Series, with a peak in abundance during Stage 3 and 4 of Series 2, and then rapidly disappearing in the Wuliuan Stage of the Miaolingian Series. We here report Oldhamia radiata from the Cambrian Series 2 Xinji Formation in western Henan of North China, associated with microbial mats, as evidenced by the presence of wrinkle structures. The distinctive patterns indicate the exploitation of microbial mats by the tracemakers of O. radiata, featuring a specific feeding strategy under the mats where possibly anoxic and sulfidic conditions prevailed during the early Cambrian.     

Ecological innovations in the Cambrian and the origins of the crown group phyla

Simulation studies of the early origins of the modern phyla in the fossil record, and the rapid diversification that led to them, show that these are inevitable outcomes of rapid and long-lasting radiations. Recent advances in Cambrian stratigraphy have revealed a more precise picture of the early bilaterian radiation taking place during the earliest Terreneuvian Series, although several ambiguities remain. The early period is dominated by various tubes and a moderately diverse trace fossil record, with the classical ‘Tommotian’ small shelly biota beginning to appear some millions of years after the base of the Cambrian at ca 541 Ma. The body fossil record of the earliest period contains a few representatives of known groups, but most of the record is of uncertain affinity. Early trace fossils can be assigned to ecdysozoans, but deuterostome and even spiralian trace and body fossils are less clearly represented. One way of explaining the relative lack of clear spiralian fossils until about 536 Ma is to assign the various lowest Cambrian tubes to various stem-group lophotrochozoans, with the implication that the groundplan of the lophotrochozoans included a U-shaped gut and a sessile habit. The implication of this view would be that the vagrant lifestyle of annelids, nemerteans and molluscs would be independently derived from such a sessile ancestor, with potentially important implications for the homology of their sensory and nervous systems.     

A cryptic record of Burgess Shale‐type diversity from the early Cambrian of Baltica

Exceptionally preserved ‘Burgess Shale‐type’ fossil assemblages from the Cambrian of Laurentia, South China and Australia record a diverse array of non‐biomineralizing organisms. During this time, the palaeocontinent Baltica was geographically isolated from these regions, and is conspicuously lacking in terms of comparable accessible early Cambrian Lagerstätten. Here we report a diverse assemblage of small carbonaceous fossils (SCFs) from the early Cambrian (Stage 4) File Haidar Formation of southeast Sweden and surrounding areas of the Baltoscandian Basin, including exceptionally preserved remains of Burgess Shale‐type metazoans and other organisms. Recovered SCFs include taxonomically resolvable ecdysozoan elements (priapulid and palaeoscolecid worms), lophotrochozoan elements (annelid chaetae and wiwaxiid sclerites), as well as ‘protoconodonts’, denticulate feeding structures, and a background of filamentous and spheroidal microbes. The annelids, wiwaxiids and priapulids are the first recorded from the Cambrian of Baltica. The File Haidar SCF assemblage is broadly comparable to those recovered from Cambrian basins in Laurentia and South China, though differences at lower taxonomic levels point to possible environmental or palaeogeographical controls on taxon ranges. These data reveal a fundamentally expanded picture of early Cambrian diversity on Baltica, and provide key insights into high‐latitude Cambrian faunas and patterns of SCF preservation. We establish three new taxa based on large populations of distinctive SCFs: Baltiscalida njorda gen. et sp. nov. (a priapulid), Baltichaeta jormunganda gen. et sp. nov. (an annelid) and Baltinema rana gen. et sp. nov. (a filamentous problematicum).     

Unlocking the early fossil record of the arthropod central nervous system

Extant panarthropods (euarthropods, onychophorans and tardigrades) are hallmarked by stunning morphological and taxonomic diversity, but their central nervous systems (CNS) are relatively conserved. The timing of divergences of the ground pattern CNS organization of the major panarthropod clades has been poorly constrained because of a scarcity of data from their early fossil record. Although the CNS has been documented in three-dimensional detail in insects from Cenozoic ambers, it is widely assumed that these tissues are too prone to decay to withstand other styles of fossilization or geologically older preservation. However, Cambrian Burgess Shale-type compressions have emerged as sources of fossilized brains and nerve cords. CNS in these Cambrian fossils are preserved as carbon films or as iron oxides/hydroxides after pyrite in association with carbon. Experiments with carcasses compacted in fine-grained sediment depict preservation of neural tissue for a more prolonged temporal window than anticipated by decay experiments in other media. CNS and compound eye characters in exceptionally preserved Cambrian fossils predict divergences of the mandibulate and chelicerate ground patterns by Cambrian Stage 3 (ca 518 Ma), a dating that is compatible with molecular estimates for these splits.     

Skimming the surface with Burgess Shale arthropod locomotion

The first arthropod trackways are described from the Middle Cambrian Burgess Shale Formation of Canada. Trace fossils, including trackways, provide a rich source of biological and ecological information, including direct evidence of behaviour not commonly available from body fossils alone. The discovery of large arthropod trackways is unique for Burgess Shale-type deposits. Trackway dimensions and the requisite number of limbs are matched with the body plan of a tegopeltid arthropod. Tegopelte, one of the rarest Burgess Shale animals, is over twice the size of all other benthic arthropods known from this locality, and only its sister taxon, Saperion, from the Lower Cambrian Chengjiang biota of China, approaches a similar size. Biomechanical trackway analysis demonstrates that tegopeltids were capable of rapidly skimming across the seafloor and, in conjunction with the identification of gut diverticulae in Tegopelte, supports previous hypotheses on the locomotory capabilities and carnivorous mode of life of such arthropods. The trackways occur in the oldest part (Kicking Horse Shale Member) of the Burgess Shale Formation, which is also known for its scarce assemblage of soft-bodied organisms, and indicate at least intermittent oxygenated bottom waters and low sedimentation rates.     

Biomechanical analyses of Cambrian euarthropod limbs reveal their effectiveness in mastication and durophagy

Durophagy arose in the Cambrian and greatly influenced the diversification of biomineralized defensive structures throughout the Phanerozoic. Spinose gnathobases on protopodites of Cambrian euarthropod limbs are considered key innovations for shell-crushing, yet few studies have demonstrated their effectiveness with biomechanical models. Here we present finite-element analysis models of two Cambrian trilobites with prominent gnathobases—Redlichia rex and Olenoides serratus—and compare these to the protopodites of the Cambrian euarthropod Sidneyia inexpectans and the modern American horseshoe crab, Limulus polyphemus. Results show that L. polyphemus, S. inexpectans and R. rex have broadly similar microstrain patterns, reflecting effective durophagous abilities. Conversely, low microstrain values across the O. serratus protopodite suggest that the elongate gnathobasic spines transferred minimal strain, implying that this species was less well-adapted to masticate hard prey. These results confirm that Cambrian euarthropods with transversely elongate protopodites bearing short, robust gnathobasic spines were likely durophages. Comparatively, taxa with shorter protopodites armed with long spines, such as O. serratus, were more likely restricted to a soft food diet. The prevalence of Cambrian gnathobase-bearing euarthropods and their various feeding specializations may have accelerated the development of complex trophic relationships within early animal ecosystems, especially the ‘arms race'' between predators and biomineralized prey.     

Small carbonaceous fossils (SCFs) from the Terreneuvian (lower Cambrian) of Baltica

We describe a new assemblage of small carbonaceous fossils (SCFs) from diagenetically minimally altered clays and siltstones of Terreneuvian age from the Lontova and Voosi formations of Estonia, Lithuania and Russia. This is the first detailed account of an SCF assemblage from the Terreneuvian and includes a number of previously undocumented Cambrian organisms. Recognizably bilaterian‐derived SCFs include abundant protoconodonts (total‐group Chaetognatha), and distinctive cuticular spines of scalidophoran worms. Alongside these metazoan remains are a range of protistan‐grade fossils, including Retiranus balticus gen. et sp. nov., a distinctive funnel‐shaped or sheet‐like problematicum characterized by terminal or marginal vesicles, and Lontohystrichosphaera grandis gen. et sp. nov., a large (100–550 μm) ornamented vesicular microfossil. Together these data offer a fundamentally enriched view of Terreneuvian life in the epicratonic seas of Baltica, from an episode where records of non‐biomineralized life are currently sparse. Even so, the recovered assemblages contain a lower diversity of metazoans than SCF biotas from younger (Stage 4) Baltic successions that represent broadly equivalent environments, echoing the diversification signal recorded in the coeval shelly and trace‐fossil records. Close comparison to the biostratigraphical signal from Fortunian small shelly fossils supports a late Fortunian age for most of the Lontova/Voosi succession, rather than a younger (wholly Stage 2) range.     

Nutritional biology: a neglected basic discipline of nutritional science

On the basis of a scientific-philosophical analysis, this paper tries to show that the approaches in current nutritional science—including its subdisciplines which focus on molecular aspects—are predominantly application-oriented. This becomes particularly evident through a number of conceptual problems characterized by the triad of ‘dearth of theoretical foundation,’ ‘particularist research questions,’ and ‘reductionist understanding of nutrition.’ The thesis presented here is that an interpretive framework based on nutritional biology is able to shed constructive light on the fundamental problems of nutritional science. In this context, the establishment of ‘nutritional biology’ as a basic discipline in research and education would be a first step toward recognizing the phenomenon of ‘nutrition’ as an oecic process as a special case of an organism–environment interaction. Modern nutritional science should be substantively grounded on ecological—and therefore systems biology as well as organismic—principles. The aim of nutritional biology, then, should be to develop near-universal ‘law statements’ in nutritional science—a task which presents a major challenge for the current science system.     

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.     

First report of Schizopholis (Lingulata,Brachiopoda) from the Tsinghsutung Formation (Cambrian Series 2, Stage 4) in Guizhou,China

Schizopholis Waagen, 1885 is a genus of linguliform brachiopod, which is known from Cambrian Stage 4 to the Wuliuan Stage of Australia, Antarctica, Pakistan and China. Recently, new material of Schizopholis was discovered from the upper part of the Tsinghsutung Formation (Cambrian Series 2, Stage 4) near Balang village, Jianhe County, Guizhou Province, China. These specimens display the oval pedicle opening in the ventral valve, a median tongue and a pair of tubercles in the dorsal valve that are characteristic features of Schizopholis napuru (Kruse, 1990). This is the first time that this species has been described from Guizhou Province, China. Previously, fossils of this species are usually preserved in carbonate deposits, but the new material documented herein preserved in mudstone of the Tsinghsutung Formation, providing new information regarding both the paleogeographic distribution and paleoecology of this species.     

A new early Cambrian bivalved euarthropod from Yunnan,China and general interspecific morphological and size variations in Cambrian hymenocarines

The hymenocarines were a major and diverse group of euarthropods featured by a bivalved head carapace and a segmented trunk comprising tergo-pleural rings notably from the Cambrian Burgess Shale-type Lagerstätten. Here we document a new ~50 mm long hymenocarine Xiazhuangocaris chenggongensis n. gen. n. sp. from the Xiazhuang fossil assemblage, a slightly younger occurrence of the Chengjiang Lagerstätte within the lower Cambrian Hongjingshao Formation (Cambrian Series 2, Stage 3) in Chenggong, Yunnan, southwestern China. X. chenggongensis is characterized by the bivalved carapace with each valve bearing a protruded anterior tip and a prominent posterior notch, which resembles the P-element of the radiodont Hurdia and is unique among the hymenocarines. The trunk consists of at least 13 tergo-pleural rings including presumably eight abdominal ones terminating in a pair of ovate caudal rami. With the body organization and morphological features closest to waptiids, X. chenggongensis represents a ‘larger’ waptiid-like hymenocarine. The morphometric analysis recognizes the family-level groupings of Cambrian hymenocarines based on the number of tergo-pleural rings and the proportion of exposed trunk length, indicating that these are useful characters for studying hymenocarine morphology and phylogeny. Cambrian hymenocarines exhibited great interspecific body size variations even between sister species of the same family, further suggesting that niche diversification happened on the family level and even among phylogenetically close genera or species during the Cambrian explosion.     

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。     

Taphonomic experiments imply a possible link between the evolution of multicellularity and the fossilization potential of soft‐bodied organisms

The reliability of evolutionary reconstructions based on the fossil record critically depends on our knowledge of the factors affecting the fossilization of soft‐bodied organisms. Despite considerable research effort, these factors are still poorly understood. In order to elucidate the main prerequisites for the preservation of soft‐bodied organisms, we conducted long‐term (1–5 years) taphonomic experiments with the model crustacean Artemia salina buried in five different sediments. The subsequent analysis of the carcasses and sediments revealed that, in our experimental settings, better preservation was associated with the fast deposition of aluminum and silicon on organic tissues. Other elements such as calcium, magnesium, and iron, which can also accumulate quickly on the carcasses, appear to be much less efficient in preventing decay. Next, we asked if the carcasses of uni‐ and multicellular organisms differ in their ability to accumulate aluminum ions on their surface. The experiments with the flagellate Euglena gracilis and the sponge Spongilla lacustris showed that aluminum ions are more readily deposited onto a multicellular body. This was further confirmed by the experiments with uni‐ and multicellular stages of the social ameba Dictyostelium discoideum. The results lead us to speculate that the evolution of cell adhesion molecules, which provide efficient cell–cell and cell–substrate binding, probably can explain the rich fossil record of soft‐bodied animals, the comparatively poor fossil record of nonskeletal unicellular eukaryotes, and the explosive emergence of the Cambrian diversity of soft‐bodied fossils.     

First evidence for Cambrian glaciation provided by sections in Avalonian New Brunswick and Ireland: Additional data for Avalon–Gondwana separation by the earliest Palaeozoic

The first evidence for Cambrian glaciation is provided by two successions on the Avalon microcontinent. The middle lowest Cambrian (middle Terreneuvian Series and Fortunian Stage–Stage 2 boundary interval) has an incised sequence boundary overlain by a fluvial lowstand facies and higher, olive green, marine mudstone on Hanford Brook, southern New Brunswick. This succession in the lower Mystery Lake Member of the Chapel Island Formation may be related to melting of an ice sheet in Avalon. The evidence for this interpretation is a muddy diamictite with outsized (up to 10 cm in diameter), Proterozoic marble and basalt clasts that penetrated overlying laminae in the marine mudstone. That eustatic rise was associated with the mudstone deposition is suggested by an approximately coeval rise that deposited sediments with Watsonella crosbyi Zone fossils 650 km away in Avalonian eastern Newfoundland. A sea-level rise within the Watsonella crosbyi Chron, at ca. 535 Ma, may correspond to a unnamed negative ¹³C excursion younger than the basal Cambrian excursion (BACE) and the ZHUCE excursion in Stage 2 of the upper Terreneuvian Series. Cambrian dropstones are now also recognized on the northern (Gander) margin of Avalon in continental slope–rise sedimentary rocks in southeast Ireland. Although their age (Early–Middle Cambrian) is poorly constrained, dropstones in the Booley Bay Formation provide additional evidence for Cambrian glaciation on the Avalon microcontinent. Besides providing the first evidence of Cambrian glaciation, these dropstone deposits emphasize that Avalon was not part of or even latitudinally close to the terminal Ediacaran–Cambrian, tropical carbonate platform successions of West Gondwana.     

寒武系小碳化石(SCFs)的研究进展及意义

所谓SCFs(Small Carbonaceous Fossils),可译为"小碳化石",其大小跨越了传统的微观与宏观界线(以肉眼可见与否为标准),是一个特定化石群体的统称。它的实质含义,是指通过氢氟酸(HF)等无机酸浸泡泥岩/页岩样品后经过滤、浓缩而获得的一类体积微小的有机碳质薄膜化石,包括后生动物残片、丝状藻/菌类碎片、疑源类等类型。在小碳化石概念被正式提出之前,国外学者曾先后用有机质壁微体化石(Organic-walled microfossils)、碳质微体化石(Carbonaceous microfossils)与布尔吉斯页岩型微体化石(Burgess Shale-type microfossils)来表述此类化石。目前在国际上,与布尔吉斯页岩型(BST)化石库和奥斯坦(Orsten)型化石库一样,小碳化石已经成为寒武纪古生物学及生物埋藏学等领域研究的一个重要的新窗口;同时,通过三类特异埋藏化石的对比研究,实现优势互补,从而能够更好地阐述早期生物的演化与多样性。     

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.     

Trace fossils from lower Cambrian Hongjingshao Formation,Yunnan, China: Taxonomy,palaeoecology, palaeoenvironment

The Hongjingshao Formation (Cambrian Series 2, Stage 3) yields abundant, complex trace fossils that have not been systematically investigated up till now. Our detailed ichnological study of the traces from this formation in Malong area, Yunnan, China, recognizes four groups: (1) simple horizontal or sub-horizontal burrows; (2) complex branched burrow systems; (3) arthropod trails and; (4) simple vertical burrows belonging to the Cruziana ichnofacies. A trophic web is reconstructed for the community in Hongjingshao Formation based on both trace and body fossils. The ichnological and sedimentological features of the formation indicate an intertidal setting. The trace fossil assemblage demonstrates that Early Cambrian organisms were able to colonize very shallow marine environments, which further supports the landward expansion of the Early Cambrian ecosystems.