New Bivalved Arthropods from the Cambrian (Series 3, Drumian Stage) of Western Hunan,South China

We report two new three-dimensionally phosphatized microfossils, Cambrolongispina reticulata gen. et sp. nov. and Cambrolongispina glabra gen. et sp. nov., from the middle Cambrian （Series 3, Drumian Stage） at Wangcun, Yongshun County, western Hunan, South China. They are bivalved arthropods, with thin, pliable, originally chitinous or chitin-calcareous shields （c. 350-517 I1 m long）. The shields are equipped with a pair of antero-dorsal spines. The spines are internally hollow, varying in length from 1/2 of to as long as the shield length in C. reticulata, and uniformly longer than the shield length in C. glabra. The spines of C. reticulata are ornamented with a longitudinal row of conical or blade-shaped denticles along the posterior edge. Cambrolongispina lacks marginal rims, valve lobation and sulci which are diagnostic of the Bradoriida sensu stricto. It also lacks a donblure/duplicature characteristic of phosphatocopids and some bradoriids. It could be related to the Monasteriidae （which may belong to Bradoriida sensu lato）, both characterized by a pair of antero-dorsal spines. Cambrolongispina may have been meiofaunai detritus feeders that lived on or within sediments. The antero-dorsal spines may have been sensory organs to detect nearby predators. The posterior denticles on these spines might have facilitated the sensory function of the spines.     

Reproductive strategy of the bradoriid arthropod Kunmingella douvillei from the Lower Cambrian Chengjiang Lagerstätte,South China

The bivalved bradoriid arthropod Kunmingella douvillei (Mansuy, 1912) is the most common species in the Lower Cambrian Chengjiang Lagerstätte. Its soft anatomy has been reported based on well-preserved specimens. However, as with other Cambrian arthropods, its reproductive behavior is poorly documented. Rare specimens of K. douvillei preserved with eggs have been reported. Our new collections confirm that the eggs were directly attached to the last three pairs of biramous appendages and verify the head–trunk boundary suggested by previous work. The brooding behavior of K. douvillei was most probably related to its small body size, as in living crustaceans. The K. douvillei female could carry 50–80 eggs ranging from 150–180 μm in diameter, possibly indicating a K-reproductive strategy; its ontogenetic and reproductive mode might have been an adaptation in response to increasing predatory pressure across the Ediacaran–Cambrian boundary. All of the three main directions of development in the life history of living marine invertebrates, namely planktotrophic larvae, lecithotrophic larvae, and brood care, most likely evolved by Stage 3 of the Cambrian.     

Addressing the Chengjiang conundrum: A palaeoecological view on the rarity of hurdiid radiodonts in this most diverse early Cambrian Lagerstätte

Over one hundred arthropod fossil species have been described from the famous Chengjiang Lagerstätte (South China, Cambrian Stage 3, ca. 518 Ma) including a diverse assemblage of radiodonts–a group containing Anomalocaris and its relatives. These iconic stem-group euarthropods include some of the largest animals of the time, and some are known from hundreds of specimens. A longstanding conundrum has been the rarity or absence of hurdiids from Cambrian Series 2 Lagerstätten like Chengjiang. This is because radiodonts are generally common in such deposits and the oldest radiodont ever discovered is a hurdiid. Furthermore, this family displays the widest geographic and temporal ranges of all radiodont families, and the highest diversity. Here we document the first hurdiid frontal appendages from Chengjiang, which display unique features within the family and may provide insights for understanding the character evolution of hurdiid appendages. The palaeoenvironmental distribution of hurdiids suggests that the rarity of hurdiids in Chengjiang may be due to a preference for deeper water environments, and the later success of this family from the Wuliuan onwards may relate to their ability to tolerate cooler water temperatures than other radiodont families. The palaeogeographical, palaeoenvironmental, and stratigraphical patterns observed in hurdiids maybe caused in part by the limited distributions of Konservat-Lagerstätten in the Cambrian as well.     

Early Cambrian epibolic gastrulation: A perspective from the Kuanchuanpu Member, Dengying Formation, Ningqiang, Shaanxi, South China

The Cambrian explosion is marked by the apparent sudden appearance of diverse metazoan skeletal fossils and an increase in the complexity of both body and trace fossils. Among them, fossil eggs and embryos from the Lower Cambrian Kuanchuanpu Member of the Dengying Formation at Ningqiang in southern China provide a unique window for investigating the ontogeny of metazoans. Gastrulation is the cell migration stage after cleavage, and can be viewed as the embryonic analog of the transition from protozoan to metazoan grades of complexity. As an example, the embryonic developmental sequences of Punctatus emeiensis is well-documented because of the stellate spines covering the body surface, that are shared with embryonic stages represented by Olivooides and their growth stages represented by P. emeiensis. Although Olivooides was considered to be a taxonomic mixture, consideration of its variation was neglected. A specific type of embryo with zigzag blastopore lips that differs from typical candidate embryos for P. emeiensis is identified; its gastrulation process is reconstructed and is comparable to the epibolic gastrulation of extant metazoans. Both the embryos and adults of this ‘type’ display radial symmetry, that suggests an affinity distinct from that of echinoderms. Moreover, the body plan of penta-radial symmetry seen widely in Early Cambrian ‘Small Shelly Fossils’ (SSF's) was probably deep rooted in the Ediacaran.     

Taxonomy and stratigraphy of Early Cretaceous species of Debrunia Masse and Fenerci-Masse (Hippuritida,Monopleuridae) of the Mediterranean region

The genus Debrunia is characterized by its myophoral organisation: the left valve posterior myophore is an erect plate, the anterior is a bulge or a crest, myophores of right valve are on shell wall, the ligament groove is inconspicuous and the cardinal apparatus tends to be shifted posteriorly. Shell habit varies from low conical to elongated cylindrical, and the shape of the left valve ranges from capuloid to flat. The taxonomic revision of Debrunia species of the Mediterranean regions reveals the presence of thirteen species, among which seven are described as new. Multiple Component Analysis and Cluster Analysis document the existence of five groups of species in correspondence with the height of the left valve and the commissural diameter. The longevity of Debrunia is high, 32 Myr, whereas its species have shorter durations which account for a significant biostratigraphic potential within the Valanginian–Albian time span. Debrunia has its FO (First occurrence) in the lower Valanginian and its LO (Last occurrence) in the Albian. Its developmental phase starts in the Hauterivian, marked by an increasing species diversity and a biogeographic expansion; and peaks in the late Barremian, followed by a collapse in diversity and a specific turnover, coeval with the “Agriopleura extinction event”. The mid-Aptian crisis is expressed by a drastic decline in species diversity, controlled by the overall restriction or demise of carbonate platforms in European regions. As elevators, Debrunia species tend to build dense monospecific aggregations and some of them possess a remarkable bioconstructional potential. The contribution of Debrunia to rudist communities is significant throughout the history of the genus.     

First skeletal remains of the giant sawfish Onchosaurus (Neoselachii,Sclerorhynchiformes) from the Upper Cretaceous of northeastern Italy

Sclerorhynchiform sawfishes are a diverse and extinct clade of elasmobranchs that is restricted to the Cretaceous. Most taxa are known only by isolated rostral spines, whereas skeletal remains are rare and have been reported from a small number of Upper Cretaceous localities. Here, we describe skeletal remains of the giant sclerorhynchiform Onchosaurus pharao for the first time, which provides new morphological information. The single specimen comes from middle-basal upper Turonian strata of the Lessini Mountains in northeastern Italy and represents the first record of this genus from Italy. The specimen consists of unidentifiable cranial remains, several diagnostic rostral spines, the rostrum with fragments of tessellated calcified cartilage, and 87 disarticulated vertebrae. The rostrum preserves the characteristic sensory system of sclerorhynchiforms. It is devoid of any lateral sockets indicating that rostral spines were attached laterally to its surface. This pattern is identical to most sclerorhynchiforms and extant pristiophoriformes implying also similar replacement patterns as in most other sclerorhynchiforms with the exception for Schizorhiza. Additionally, the bases of two longitudinally arranged rows of ventral rostral spines are identifiable concurring with patterns seen in Sclerorhynchus. The axial skeleton is partly preserved. Re-arranging the disarticulated vertebrae according to their life position in combination with measures of the size and thickness of preserved vertebral centra, and the ratio rostrum length/body size depending on the number of vertebral centra indicate that the specimen was ca. 450 cm long. Growth rings in the vertebral centra show that the specimen was about four years old and thus probably not yet fully sexual mature when it died. This age assumption corresponds well with the calculated size when compared with complete skeletons of extinct sclerorhynchiforms and extant pristiforms. The size of the specimen and its occurrence in hemipelagic rocks corroborates previous assumptions that this sclerorhynchiform was a large and pelagic sawfish.     

Cambrian stratigraphy of Kashmir and its boundary problems

The biostratigraphy of the Cambrian of Kashmir based on trilobite assemblage zones and trace fossils has been attempted. Faunal gaps occur in lower part of Lower Cambrian, upper Lower Cambrian, lower Middle Cambrian and upper part of Upper Cambrian. In the Lower Cambrian Cruziana—Rusophycus and Redlichia Zones, in the Middle Cambrian Solenopleura—Tonkinella, Tonkinella—Anomocare, Anomocare—Bailiella and Bolaspidella Zones, and in the Upper Cambrian Chuangia and Dikelocephalites Zones are recognized. The position of intrasystem boundaries is also discussed. While the various taxa have lithological preferences and are not necessarily found in a continuous sequence, an attempt has been made to interpret the ranges of different genera. The affinities of the fauna with that of other Cambrian basins of the world are discussed. It is concluded that the bulk of the fauna is of a cosmopolitan nature.     

Lower Cambrian helcionelloid macromolluscs from South Australia

Early Cambrian univalve molluscs are predominantly represented by microscopic forms (body length of 1–3 mm), preserved mainly as phosphatised internal moulds with limited definable features. Macromolluscs (≥ 5 mm) are generally rare, occur in low abundance and are poorly preserved, often lacking apical features and ornament which hinders taxonomic assessment. New and previously undescribed material from lower Cambrian Hawker Group carbonates of the Flinders Ranges in South Australia includes four new taxa, Minastirithella silivreni gen. et sp. nov., Galeacalvus coronarius gen. et sp. nov., Helcionella histosia sp. nov., and Ilsanella enallaxa sp. nov. Three-dimensional silicified shell material preserved with intact apices offers new insight into protoconch morphology, growth habit (isometric vs. allometric) and developmental mode. This material supports previous suggestions that some micromollusc taxa may in fact be early ontogenetic stages (juveniles) of larger macroscopic taxa; such that the millimetric size range of helcionelloids conforms to the dimensions of earliest apical portion in some macromolluscs documented herein. However, taphonomic limitations associated with phosphatisation show that the morphology (especially height vs. width) of the apex can greatly influence the probability of steinkern formation and preservation potential for both micro- and macro-scale helcionelloids. Artificial inflation in the appearance of millimetric helcionelloids with an optimal morphology for phosphatisation is thus directly linked to taphonomic biases in the fossil record.     

Dimorphism of Bivalved Arthropod Branchiocaris? Yunnanensis from the Early Cambrian Chengjiang Biota, South China

The bivalved arthropod Branchiocaris? yunnanensis Hou, 1987 is redescribed on the basis of new fossil material from the early Cambrian Chengjiang biota, South China. In total, 81 specimens have been examined. The carapace comprises two identical valves, each valve are sub-circular in outline, ranging from 24 mm to 58 mm in length, and from 15 mm to 46 mm in height. The dorsal margin is straight and bears two short cardinal spines. The valve surface is either smooth or ornamented with polygonal reticular structures, which may result from different preservation conditions. Two morphotypes have been recognized based on the presence or absence of a subdorsal swelling of the valve, which is convex dorsolaterally and extends beyond the dorsal margin when it appears. Statistic and ontogenetic analyses show both morphotypes grow isometrically and have the same growth trajectory. Therefore, the two morphotypes are interpreted as dimorphism within the same species. Our results are helpful for investigating the taxonomy and ontogeny of arthropod fossils and suggest that dimorphism might be fairly common in these early arthropods.     

The southern border of the Guaporé Shield in western Brazil and Bolivia: An interpretation of its geologic evolution

The geology of the southeastern and southwestern margins of the Guaporé Shield is described. The Shield rocks comprise high-grade granulite and low-grade greenschist sequences. These rocks are poorly known. Oldest recorded ages include Transamazonian datings (± 2 Ma). The margins of the shield show the development of a mobile belt formed by rocks of Upper Proterozoic to Cambrian age. Within this mobile belt, the Cuiabá Group of uncertain (but possibly Upper Proterozoic) age outcrops to the east and shows tectonic contacts with younger rocks that span the Vendian (Upper Proterozoic) to Cambrian age. The rocks of Vendian to Cambrian age form three isolated elongated basins which show evidence of fault control. The axes of these basins are equally separated by angles of ～ 120° and meet at a triple junction near Corumbá. The stratigraphy of the Vendian to Cambrian basins is correlated from Bolivia to Brazil, based on tectonic continuity, sedimentological similarity and fossil content. The central location at Corumbá shows instability since the Middle Proterozoic when important igneous activity is recorded at Rincon del Tigre (Bolivia) and in Brazil. This igneous activity is interpreted as being related to hot-spot activity in an intercratonic setting from which the younger Upper Proterozoic basins originated.     

Osteoderms of Montealtosuchus arrudacamposi (Crocodyliformes,Peirosauridae) from the Turonian-Santonian (Upper Cretaceous) of Bauru Basin,Brazil

The skin of Crocodyliformes is characterized by osteoderms of various sizes and shapes. It plays roles in defence, thermoregulation, sexual attraction, calcium reserves, and locomotion. This study presents the morphometric characteristics of osteoderms preserved in the nuchal, dorsal, ventral and appendicular shield of Montealtosuchus arrudacamposi, a Peirosauridae crocodyliform from the Turonian-Santonian (Upper Cretaceous) of Adamantina Formation, Bauru Basin, Brazil. The results of the analysis show that its dermal shield had protective and thermoregulatory functions and a well-defined mechanical function allowing terrestrial locomotion and enabling good agility and capacity for movement.     

Morphology and ontogeny of Hunanocephalus ovalis (trilobite) from the Cambrian of South China

The juvenile morphology and ontogeny of the Cheiruroidid trilobite Hunanocephalus ovalis Lee, 1963 from the lower Cambrian Shuijingtuo Formation in Hubei Province, South China is presented. The new material comprises a relatively complete meraspid ontogenetic series (degree 0 to 10), which reveals more details on their morphological changes such as the contraction and disappearance of the pronounced posteromedial notch in the pygidium and the addition of the trunk segments, which are all documented for the first time and can also be used as developmental markers defining their ontogenetic phases. The trunk segmentation schedule of H. ovalis is also discussed, which is similar to the other early Cambrian oryctocephalid trilobites, i.e. as the boundary between the thorax and pygidium migrated posteriorly there is no change in the trunk segment number; the processes of liberation of the thoracic segment and segment insertion into the pygidium are separated from one another, implying that the control of trunk exoskeletal segment appearance and articulation might be decoupled in these trilobites.     

Sur la présence d'une série molassique (de type série pourprée) au Sud-Est de l'Ahaggar (In Guezzam,Ahaggar, Algérie)Presence of molassic series (‘Série pourprée’ type) in the Southeast of the Ahaggar (In Guezzam,Ahaggar, Algeria)

In the western Ahaggar shield, the transition between the Precambrian and the Ordovician units is characterised by thick volcano-sedimentary series (‘Série pourprée’ of the Ahnet). This series, in part of Cambrian age, results from the demolition of the Panafrican belt. Similar series were known in grabens located between the West African craton and the Ahaggar, from the ‘Adrar des Iforas’, in the south, to the Ougarta belt, in the north. We describe in this study a new formation identical to the ‘Série pourprée’ of the Ahnet, but cropping out in the far Southeast of the Ahaggar (In Guezzam). This new datum improves the Panafrican belt configuration. To cite this article: H. Djellit et al., C. R. Geoscience 334 (2002) 789–794.     

Systematics, biometry of the species Opsidiscus from the Middle Cambrian succession of the Spiti basin, India

The Middle Cambrian succession of the Parahio Valley in general and Debsakhad section in particular is dominated by trilobites. Genus Opsidiscus, along with other trilobite forms dominates the lower Middle Cambrian succession. In the present study two new species of Opsidiscus, Opsidiscus wadiai and Opsidiscus srikantiai are reported from the lower Middle Cambrian succession of Debsakhad section. The species were differentiated on their morphological features and their multivariate analysis. The multivariate analysis applied here is used as a supplement method to qualitative analysis in order to differentiate between the cranidial characters of Opsidiscus. The qualitative study of each species studied individually shows more or less close affiliation as is observed by different quantitative methods. The presence of Opsidiscus has a great stratigraphic significance in this region, as they first appear from informal Stage 5 of Series 3 and goes up to the Drumian Stage of Series 3 of the Cambrian System in the Debsakhad section, which helps to correlate this section with other well known sections of the Middle Cambrian.     

Mesoprosopon triasinum from the Triassic of Austria revisited: The oldest eumalacostracan larva known to date and its significance for interpreting fossil cycloids

The enigmatic Mesoprosopon triasinum from the Triassic Hallstatt Limestone of Austria, which was once considered to be either a representative of Brachyura or Cycloidea, is re-interpreted herein as a eumalacostracan larva and is the oldest of its type known to date. It shows a mixture of characters that are typical of the zoea stages of certain meiuran ingroups (e.g., Hippidae) or of erichthus-type larvae of stomatopods. Four long spines evidently provided additional buoyancy to counteract the comparatively heavy load of a calcified shield. Additionally, a distinct ventral gape might imply that the animal was able to enrol into a tight ball. Our recognition of specimens of M. triasinum as larval stages, rather than adults, may have a major impact on the re-study of some still poorly known cycloids. In the future, the term “mesoprosopon” may be salvaged as the name of this type of larva.     

Stable isotope(δ~(13)C_(ker),δ~(13)C_(carb),δ~(18)O_(carb)) distribution along a Cambrian outcrop section in the eastern Tarim Basin,NW China and its geochemical significance

This study investigated the geochemical features of the lower Paleozoic strata of Yaerdang Mountain outcrop along with the core samples from well TD2∈ in the eastern Tarim Basin,NW China.The total organic carbon abundance,hydrocarbon-generating precursor biospecies,and stable isotope ratios of organics and carbonate(δ~(13)C_(ker),δ~(13)C_(carb) and δ~(18)O_(carb)) were comprehensively studied for their possible correlative constraints during sedimentary evolution.The results revealed that the δ~(13)C_(ker)(VPDB) of Cambrian kerogens along the outcrop section varied from-34.6‰ to-28.4‰,indicating an increasing tendency from the lower Cambrian to the upper Cambrian.This was on the whole accompanied by the variation in the δ~(13)C_(carb) and δ~(18)O_(carb) along the profile,which might be associated with the changes in the sea level and also in the compositional variation of benthic and planktonic biomass.The large variation in the stable carbon isotope ratios up to 6‰ along the outcrop section reflected the heterogeneity of the Cambrian source rocks from the eastern Tarim Basin.Hence,the ~(13)C-enriched crude oils from well TD2∈might have been derived from a localized stratum of Cambrian source rocks.The results from this study showed the possibility of multiple source kitchens in the Cambrian-lower Ordovician portion of Tarim Basin.     

The oldest bivalved arthropods from the early Cambrian of East Gondwana: Systematics,biostratigraphy and biogeography

The oldest bradoriid fauna from Australia, occurring in the lower Cambrian Ajax and Wirrapowie limestones of the Flinders Ranges, South Australia consists of eleven taxa, including one new genus and species, Quadricona madonnae gen. et sp. nov. and two new species, Liangshanella circumbolina sp. nov. and Zepaera jagoi sp. nov. In the Ajax Limestone, Liangshanella circumbolina sp. nov. occurs c. 20 m below the FAD of the zonal trilobite Abadiella huoi. This pre-trilobitic occurrence represents the oldest bivalved arthropod hitherto known from East Gondwana and suggests a lower Cambrian (Series 2, Stage 3) age for the assemblage. The recognition of distinct bradoriid assemblages associated with the Abadiella huoi (Atdabanian), Pararaia tatei, P. bunyerooensis and P. janeae (all Botoman) trilobite biozones in South Australia indicates great potential for future regional biostratigraphic correlation. Quantitative biogeographic analysis including new taxonomic data from the lower Cambrian of South Australia, highlights the strong endemism displayed by early Cambrian bradoriid communities and strengthens the close faunal affinities with South China and Antarctica.     

Quantitative taxonomy and biostratigraphy of Middle Cambrian trilobites from Montana and Wyoming

This study was made to revise the taxonomy and biostratigraphy of Middle Cambrian trilobites from Montana and Wyoming. Both numerical and conventional analyses were made of the taxonomic and biostratigraphic data. Numerical taxonomy was performed on a sample of 210 OTU's (Operational Taxonomic Units) using 66 characters which were determined on cranidia of Middle Cambrian ptychopariid trilobites. These characters consisted of linear measurements which were transformed to indices proportional to glabellar length, quantitative-qualitative shape or angle measurements, and multistate attributes. The similarity coefficients calculated were average taxonomic distances and Pearson's product-moment correlation coefficient. The matrices of similarity coefficients were clustered by single-linkage and unweighted pair-group algorithms. The coordinates of OTU's in a three-dimensional A-space defined by the first three principal axes also were calculated. A phenogram of the distance matrix clustered by UPGMA (which yielded the highest cophenetic correlation coefficient) and a three-dimensional pin-and-ball model were used to interpret the trilobite taxonomy. Insufficiently clear clustering in both representations of taxonomic structure necessitated also a conventional taxonomic study guided by the numerical phenetics. Numerical biostratigraphy was performed with 79 collections as OTU's and 43 trilobite genera as characters. Jaccard coefficients were used as similarity coefficients, and the results were clustered by UPGMA. The resultant phenogram was readily interpretable, and seven faunal assemblage zones (Albertella, Glossopleura, Ehmaniella, Bolaspis-Glyphaspis, Ehmania, Parehmania, and Bolaspidella)are defined for the Middle Cambrian in the study area. Two of the zones (Ehmaniellaand Parehmania)are new and replace part of the previously defined “Bathyuriscus-Elrathina”zone. Bolaspis-Glyphaspis, Ehmania,and Parehmaniazones may have only regional validity.     

Additional late Middle Cambrian trilobites from Karsha Formation (Haimanta Group) Zanskar region of Zanskar-Spiti basin, Northwest Himalaya

Additional latest Middle Cambrian polymerid trilobites are recorded from the Teta Member (Karsha Formation) along the Tangze-Yogma-Kuru section, Kurgiakh Valley, southeastern Zanskar, Northwest Himalaya. It includes record of a new genus Pianspis Saito & Sakakuru, 1936 of family Lisianiidae and others associated fauna comprises of Neoanomocarella asiatica (Lisianiidae), Parablackwelderia sp., (Damesellidae) and two undetermined pygidium and one unidentified cranidia. All these specimens are collected from the level representing the Lejopyge acantha Zone of Guzhangian Stage (Cambrian Series 3) of the Middle Cambrian. The fauna is mainly correlated with the South China, North Korea, Kazakhastan, Uzbekistan, Siberia and Australia.     

Chlorine stable isotope distribution of Michigan Basin formation waters

The Cl isotope ratio, mass³⁷Cl/³⁵Cl, was measured on 22 formation waters from Mississippian, Devonian, Silurian, Ordovician and Cambrian strata in the Michigan Basin. Because of its resistance to fractionation, the ratio was used to find evidence of mixing of formation waters within the Michigan Basin and between the Canadian Shield and the basin. The δ³⁷Cl composition of waters decreased from +0.05 to −0.55 (per mil difference from SMOC, precision of 0.16) with age of the strata among Devonian, Silurian, Ordovician and Cambrian samples from the basin margin. Mississippian samples from the middle of the basin were isotopically heaviest at +0.1. Ordovician samples, also from mid-basin, were isotopically lightest at −1.2. On plots of δ³⁷ Cl vs Cl/Br and δ³⁷ Cl vs⁸⁷Sr/⁸⁶Sr samples at the basin margin trend toward enrichment in³⁵Cl and⁸⁷Sr and increasing Cl/Br suggesting interformational mixing of the waters. On a δ³⁷ ClCl/Br plot, three samples not on this trend and tending toward high Cl/Br may reflect evaporite dissolution. Canadian Shield Waters were plotted with Michigan Basin waters on the graphs of δ³⁷ Cl vs Ca/Cl and δ³⁷ Cl vs K/Cl. On both plots data fall along linear trends of³⁵Cl depletion with Ca/Cl increase and with K/Cl decrease. Ordovician waters from the middle of the basin and shield waters are end members on the plots. The results suggest that despite water-rock interactions, δ³⁷ Cl data may be useful in studies of mixing relations in formation waters.