Problematic cap-shaped fossils from the Lower Cambrian of North-East Greenland

Three species ofOcruranus Liu, 1979 are described from the Bastion Formation of North-East Greenland, of late Early Cambrian (middle Dyeran of North American usage) age, representing the youngest record of a genus originally described from the earliest Cambrian Meishucunian Stage of China. An accompanying species, tentatively assigned toXianfengella He &Yang, 1982, seems also to be present in South Australia in strata of late Early Cambrian (Botoman of Siberian usage) age, although also this genus was described originally from the Meishucunian.Ocruranus andXianfengella from China have been interpreted as possibly parts of coeloscleritophoran scleritomes, perhaps halkieriids, rather than individual molluscan shells. Their shell form is not typical of helcionelloids which otherwise dominate the Early Cambrian molluscan record, but conclusive evidence of affinity is not forthcoming from the Greenland records. New taxa:Ocruranus septentrionalis n. sp. andOcruranus tunuensis n. sp.      

Moulting in the lobopodian Onychodictyon from the lower Cambrian of Greenland

A number of lobopodian taxa from the Cambrian display pairs of sclerotized plates symmetrically positioned along the dorsum of the animal, predominantly above the walking appendages. Most genera were described from complete body fossils exquisitely preserved in the famous Cambrian Lagerstätten, but lobopodian phosphatized plates are found worldwide as typical components of Cambrian small shelly fossil assemblages (SSF). Details regarding intraspecific and ontogenetic variation in lobopod plates are elusive, and the lack of details of ornamentation in Lagerstätte specimens does not minimize the problem. We document here an assemblage of well‐preserved isolated plates of Onychodictyon sp. from the Lower Cambrian (Cambrian Series 2, Stage 4) of North Greenland. Two specimens exhibit perfectly conjoined plates from successive moults. Details of ornamentation and the outline and profile of the fixed plates are identical, but width and length of the underlying plate are 24% larger. These specimens boost the body of evidence that lobopodians moulted but also show that plate outline and ornamentation did not vary during ontogeny.     

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.     

A New Bivalved Arthropod from the Early Cambrian Sirius Passet Fauna, North Greenland

A new bivalved arthropod is described from the Lower Cambrian (?Upper Atdabanian) Buen Formation of North Greenland. Pauloterminus spinodorsalis gen. et sp. nov. possesses a bivalved carapace that covers the head, which has a single pair of antennae, and anteriormost thorax. No mouthparts are visible. The five‐segmented abdomen was limbless and terminated in a telson plus a pair of large, lobate uropods. A suite of at least six biramous thoracic limbs are present: the short endopods are made up of small, serial podomeres, while the exopods are lobate and may have functioned as gills as well as in swimming. Partially infilled guts are occasionally visible, suggesting that this animal may have been a sediment feeder. It is compared to other Cambrian bivalved arthropods, especially the waptiids Chuandianella ovata from the Lower Cambrian Chengjiang fauna (China) and Waptia fieldensis from the Middle Cambrian Burgess Shale (British Columbia). Of these three animals, the Greenland and Chinese taxa appear to be the most closely related. P. spinodorsalis possesses many typical arthropod features, but it also demonstrates more primitive characters that are more reminiscent of the lobopodians.     

Isoxys (Arthropoda) with preserved soft anatomy from the Sirius Passet Lagerstätte,lower Cambrian of North Greenland

Stein, M., Peel, J.S., Siveter, D.J. & Williams, M. 2009: Isoxys (Arthropoda) with preserved soft anatomy from the Sirius Passet Lagerstätte, lower Cambrian of North Greenland. Lethaia, Vol. 43, pp. 258–265. Isoxys volucris is the most commonly occurring species in the lower Cambrian Sirius Passet Lagerstätte of North Greenland. Newly identified material allows a first, limited, account of the ventral morphology of this species, hitherto known only by the morphology of its shield. The antennula is large and robust, composed of about seven articles armed with spines, and was probably not sensorial. The postantennular limbs are serially similar, biramous with a large paddle‐shaped exopod fringed with setae. It is possible that the animal possessed a furca. The inner lamella, lining the ventral surface of the shield is recognised in Isoxys for the first time. Comparisons with other congeneric species of which aspects of the ventral morphology are known, show similarities with Isoxys auritus from China, reconsidered here, but indicate differences in antennular morphology with other species as currently understood. □Cambrian, Greenland, Isoxys, soft anatomy, Sirius Passet, palaeoecology.     

A crown‐group demosponge from the early Cambrian Sirius Passet Biota,North Greenland

Calibration of the divergence times of sponge lineages and understanding of their phylogenetic history are hampered by the difficulty in recognizing crown versus stem groups in the fossil record. A new specimen from the lower Cambrian (Series 2, Stage 3; approximately 515 Ma) Sirius Passet Biota of North Greenland has yielded a diagnostic spicule assemblage of the extant demosponge lineages Haploscleromorpha and/or Heteroscleromorpha. The specimen has disarticulated approximately in situ, but represents an individual sponge that possessed monaxon spicules combined with a range of slightly smaller sigma, toxa and unique spiral morphologies. The combination of spicule forms, together with their relatively large size, suggests that the sponge represents the stem lineage of Haploscleromorpha + Heteroscleromorpha. This is the first crown‐group demosponge described from the early Cambrian and provides the most reliable calibration point currently available for phylogenetic studies.     

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.     

The Cambrian radiation of shelly fossils

One of the most significant events in the history of the organic world was the acquisition by animals of the ability to build a skeleton. This is of special interest because the overwhelming majority of known major groups (phyla) acquired that ability during a very short period (five to seven million years), early in the Cambrian. Recent fossil finds, especially in northern Asia, are adding much detail to our knowledge of this period.     

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.     

Trace fossils of Cambrian aglaspidid arthropods

A new trace fossil, Raaschichnus gundersoni , from the Upper Cambrian St. Lawrence Formation of Wisconsin, was produced by an aglaspidid arthropod. The rusophyciform trace, which occurs singly and in series, is distinctive in showing marks left by the tail-spine. Other trace fossils previously considered to have been made by aglaspidids were probably excavated by animals lacking a tail-spine and with appendage morphologies very different from the aglaspidids of Wisconsin.     

The problematic microfossil Utahphospha from the Upper Cambrian of the western United States

Phosphatic conical microfossils from Upper Cambrian (Franconian) limestones of the Orr Formation in western Utah are described as Utahphospha sequina n.g., n.sp. The genus is also known from other localities in Utah and Nevada. The fossil has a distinct sequin-like external sculpture which has been formed by outward accretion. A hole at the tip originated by abrasion from within. The fossils are interpreted as hatched cysts of an unknown organism.     

Deciphering the early evolution of echinoderms with Cambrian fossils

Echinoderms are a major group of invertebrate deuterostomes that have been an important component of marine ecosystems throughout the Phanerozoic. Their fossil record extends back to the Cambrian, when several disparate groups appear in different palaeocontinents at about the same time. Many of these early forms exhibit character combinations that differ radically from extant taxa, and thus their anatomy and phylogeny have long been controversial. Deciphering the earliest evolution of echinoderms therefore requires a detailed understanding of the morphology of Cambrian fossils, as well as the selection of an appropriate root and the identification of homologies for use in phylogenetic analysis. Based on the sister‐group relationships and ontogeny of modern species and new fossil discoveries, we now know that the first echinoderms were bilaterally symmetrical, represented in the fossil record by Ctenoimbricata and some early ctenocystoids. The next branch in echinoderm phylogeny is represented by the asymmetrical cinctans and solutes, with an echinoderm‐type ambulacral system originating in the more crownward of these groups (solutes). The first radial echinoderms are the helicoplacoids, which possess a triradial body plan with three ambulacra radiating from a lateral mouth. Helicocystoids represent the first pentaradial echinoderms and have the mouth facing upwards with five radiating recumbent ambulacra. Pentaradial echinoderms diversified rapidly from the beginning of their history, and the most significant differences between groups are recorded in the construction of the oral area and ambulacra, as well as the nature of their feeding appendages. Taken together, this provides a clear narrative of the early evolution of the echinoderm body plan.     

Mobergellans (Problematica) from the Cambrian of Greenland, Siberia and Kazakhstan

Mobergellans are disc-shaped problematic fossils of phosphatic composition bearing prominent muscle attachment structures on the presumed internal surface. They are found exclusively in sediments of Early Cambrian age, mainly from Laurentia, Baltica and Siberia. Type material ofDiscinella brastadi Poulsen, 1932 from North-East Greenland is re-studied and synonomised withDiscinella micans (Billings, 1871). Specimens from Siberia described asD. brastadi are re-assigned toMobergella hexactina n. sp., and specimens from Siberia described asMobergella radiolata Bengtson, 1968 are re-assigned toM. sibirica n. sp.Aktugaia triangula Mis-Sarzhevsky, 1976 from Kazakhstan is redescribed, and its relationship to other mobergellans is discussed.      

The Ocruranus–Eohalobia group of small shelly fossils from the Lower Cambrian of Yunnan

Lower Cambrian phosphorite samples from Maotianshan, Yunnan Province, South China, yielded a variety of small shelly fossils. Sclerites of the second pre-trilobite faunal assemblage of Meishucun, the Paragbborilus-Siphogonuchites–Lapworthella association of Nemakit–Daldynian to Tommotían age, were dominant. They comprise previously unknown spine-bearing specimens of Ocruranus finial Liu, 1979, and new morphotypes of Eohalobia diandongensis Jiang, 1982. Well-preserved sclerites reveal a laminated, composite fine structure with marginal, incremental growth. Each lamella consists of radial series of minute growth elements. The shells are phosphatically preserved, but micropetrographic evidence from drill-core samples indicates calcium carbonate as primary shell material. In the light of this new evidence, Ocruranus–Eohalobia group fossils are interpreted as parts of coeloscleritophoran scleritomes.     

Vetulicolians from the Lower Cambrian Sirius Passet Lagerstätte,North Greenland,and the polarity of morphological characters in basal deuterostomes

Abstract: Vetulicolians are problematic Cambrian fossils with a debated phylogenetic history. Here, we describe two vetulicolian specimens from the Lower Cambrian Sirius Passet locality in North Greenland. One of the specimens is assigned to Ooedigera peeli gen. et sp. nov, whereas the other is retained under open nomenclature. The mode of tail flexibility has been debated in the literature, and we argue here that the tail normally flexed laterally to generate power strokes rather than dorsoventrally. The phylogenetic relationships of vetulicolians are discussed in the light of current knowledge of deuterostome phylogeny and morphology, and it is concluded that the best hypothesis on currently available evidence is that vetulicolians are a clade or paraphyletic assemblage of stem‐Deuterostomata. The presence of a voluminous, sediment‐filled anterior chamber suggests that the atrium may be a synapomorphy of deuterostomes.     

Anabarella australis (Mollusca, Helcionelloida) from the Lower Cambrian of Greenland

The Early Cambrian helcionelloid mollusc Anabarella australis is described from North-East Greenland, representing the second occurrence of the species outside of Australia. Other Australian molluscs of this age are known from many localities including North China, Siberia, Altai, Transbaikalia, southern Kazakhstan, Mongolia, eastern Germany and Spain. These records, supported now by A. australis, demonstrate the close proximity of continents in the Early Cambrian.     

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.     

贵州织金寒武系戈仲伍组钉形类骨片化石研究*

钉形类(nail-shaped)骨片化石是华南寒武纪早期地层中常见的一类具矿化骨骼的小壳化石。本文系统研究了贵州织金戈仲伍剖面寒武系戈仲伍组的钉形类骨片化石, 发现织金地区钉形类化石主要以织金钉类和疑似的寒武钉类为主, 共4属9种。相较于前人研究, 本文新发现了织金钉类: Zhijinites tumourifomis、Z. lubricus、Z. clavus, 和疑似的寒武钉类: ?Deiradoclavus sp.、?Cambroclavus sp.。排除埋藏学假象后, 分析了钉形类不同化石属种保存方式及形态结构, 发现Zhijinites的多数种都存在盘体顶面凹陷结构, 并且Zhijinites部分种的锥体内壁有收缩现象或具有十字交错纤维结构, 这些发现能较好地揭示出Zhijinites各形态种间的相似性和差异性。分析钉形化石时空分布特点发现, 目前钉形类骨片化石最早在织金地区有化石记录, 之后在扬子地台内部辐射扩散, 并几乎同时辐射扩散至邻近陆块。此外, 本文讨论的疑似寒武钉类化石时代可下延至寒武纪幸运期, 可能是目前该类化石在全球的最早记录, 为探索寒武钉类化石的系统演化和地层学意义提供了新信息。