A contribution to sedentary polychaete phylogeny using 18S rRNA sequence data

The phylogenetic position of Annelida as well as its ingroup relationships are a matter of ongoing debate. A molecular phylogenetic study of sedentary polychaete relationships was conducted based on 70 sequences of 18S rRNA, including unpublished sequences of 18 polychaete species. The data set was analysed with maximum parsimony and maximum likelihood methods. Clade robustness was estimated by parsimony-bootstrapping and jackknifing, decay index, and clade support, as well as a posteriori probability tests using Bayesian inference. Irrespective of the applied method, some traditional sedentary polychaete taxa, such as Cirratulidae, Opheliidae, Orbiniidae, Siboglinidae and Spionidae, were recovered by our phylogenetic reconstruction. A close relationship between Orbiniidae and Questa received a particularly strong support. Echiura appears to be a polychaete ingroup taxon which is closely related to Dasybranchus (Capitellidae). As in previous molecular analyses, no support was found for the monophyly of Annelida nor for that of Polychaeta. However, we suggest that an increase in taxon sampling may yield additional resolution in the reconstruction of polychaete ingroup phylogeny, although the difficulties in reconstructing the basal phylogenetic relationships within Annelida may be due to their rapid radiation.     

Evidence for a coelomic maturation factor controlling oocyte maturation in the polychaete Arenicola marina (L.)

Summary

Previous studies on Arenicola marina suggested that oocyte maturation was induced by a single maturation hormone from the prostomium. This maturation hormone was thought to act directly on the oocyte (Meijer and Durchon, 1977), A recently described species, Arenicola defodiens (Cadman and Nelson-Smith, 1993), morphologically very similar to A. marina, has been found at the sampling sites described by Meijer and Durchon (1977). Results presented here from studies on British populations of Arenicola marina show that in this species, oocyte maturation is controlled by two hormonal steps. The first step involves the prostomial maturation hormone. The second step depends on a maturation inducing substance in the coelomic fluid. We will refer to this as the coelomic maturation factor (CMF). A reliable in vitro assay for oocyte maturation in the lugworm Arenicola marina has been adopted. It utilizes fluorescence staining of the chromosome material with DNA labelling dyes (Hoechst 33342 and 33258). Maturation of oocytes in A. marina involves germinal vesicle breakdown (GVBD). This is accompanied by the movement of chromosomes from late prophase to metaphase of meiosis I and chromosome condensation. The chromosomes are stained brightly by the dyes and their relative positions can be easily identified so that mature and immature eggs can be distinguished by the differences in chromosome position and form. The development of the in vitro fluorescence assay has enabled us to demonstrate that there are two endocrine steps involved in the induction of oocyte maturation. We have begun the characterization of CMF, and data show this to be a thermolabile molecule with a molecular mass greater than 10 kd.     

Biochemical systematics of two sibling species of polychaete annelids: Ophelia bicornis and O. radiata

Biochemical analysis of polychaete annelids from the English Channel and the Atlantic Coast has shown that they represent two genetically isolated species, Ophelia bicomis and O. radiata, characterized by different alleles at four loci and a mean number of 15 and 14 pairs of gills, respectively. Sampling of Mediterranean annelids was insufficient to determine whether or not the population studied belongs to the O. radiata species.     

Infaunal augurs of the Cambrian explosion: An Ediacaran trace fossil assemblage from Nevada,USA

Morphologically complex trace fossils, recording the infaunal activities of bilaterian animals, are common in Phanerozoic successions but rare in the Ediacaran fossil record. Here, we describe a trace fossil assemblage from the lower Dunfee Member of the Deep Spring Formation at Mount Dunfee (Nevada, USA), over 500 m below the Ediacaran–Cambrian boundary. Although millimetric in scale and largely not fabric‐disruptive, the Dunfee assemblage includes complex and sediment‐penetrative trace fossil morphologies that are characteristic of Cambrian deposits. The Dunfee assemblage records one of the oldest documented instances of sediment‐penetrative infaunalization, corroborating previous molecular, ichnologic, and paleoecological data suggesting that crown‐group bilaterians and bilaterian‐style ecologies were present in late Ediacaran shallow marine ecosystems. Moreover, Dunfee trace fossils co‐occur with classic upper Ediacaran tubular body fossils in multiple horizons, indicating that Ediacaran infauna and epifauna coexisted and likely formed stable ecosystems.     

Cambrian stem-group annelids and a metameric origin of the annelid head

The oldest fossil annelids come from the Early Cambrian Sirius Passet and Guanshan biotas and Middle Cambrian Burgess Shale. While these are among the best preserved polychaete fossils, their relationship to living taxa is contentious, having been interpreted either as members of extant clades or as a grade outside the crown group. New morphological observations from five Cambrian species include the oldest polychaete with head appendages, a new specimen of Pygocirrus from Sirius Passet, and an undescribed form from the Burgess Shale. We propose that the palps of Canadia are on an anterior segment bearing neuropodia and that the head of Phragmochaeta is formed of a segment bearing biramous parapodia and chaetae. The unusual anatomy of these taxa suggests that the head is not differentiated into a prostomium and peristomium, that palps are derived from a modified parapodium and that the annelid head was originally a parapodium-bearing segment. Canadia, Phragmochaeta and the Marble Canyon annelid share the presence of protective notochaetae, interpreted as a primitive character state subsequently lost in Pygocirrus and Burgessochaeta, in which the head is clearly differentiated from the trunk.     

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.     

Paleoecology of a large Early Cambrian bioturbator

The Lower Cambrian Poleta Formation in the White-Inyo Mountains of eastern California contains well-preserved and laterally extensive exposures of the large looping and meandering trace fossil Taphrhelminthopsis nelsoni n.isp. Such traces are typical features on upper bed surfaces of Lower Cambrian shallow marine sandstones and occur with Ediacaran fossils at other localities. Morphologic, sedimentologic and goniogram analyses suggest that the inferred tracemaker was a large soft-bodied echinozoan- or mollusc-grade animal with a volume greater than 14 cm³ that actively grazed or ingested sediment at the sediment-water interface. Although portions of these traces appear to reflect relatively 'complex' behavior, looping patterns are not periodic as expected for a systematic foraging strategy. T. nelsoni traces are patchy in distribution and commonly associated with suspect-microbial features, suggesting that tracemakers may have been targeting microbial-based or related concentrations of food resources. Such behavioral patterns are typical of shallow late Neoproterozoic-early Cambrian settings, and like suspect-microbial structures are later restricted to deep marine or stressed settings.     

Early Cambrian Teichichnus-dominated ichnofabrics and palaeoenvironmental analysis of the Caerfai Group, Southwest Wales, UK

The early Cambrian Caerfai Group of Pembrokeshire in Southwest Wales yields an intensely bioturbated Teichichnus-dominated ichnofabric, and provides evidence for the early evolution of infaunal organisms. Three units of the Caerfai Group comprise the focus of this study in sections exposed at Caerfai Bay. In the St. Non's Sandstone few primary sedimentary structures are observed, bedding being predominantly massive with subordinate parallel and current ripple cross-lamination. The topmost St. Non's Sandstone contains common soft-sediment deformation phenomena, and collectively deposition is believed to have taken place in the proximal subaqueous reaches of a steep-fronted delta system. Teichichnus is the dominant trace, with subordinate Planolites, Palaeophycus and possible Rhizocorallium.The overlying Caerfai Bay Shales were deposited from mass flows with evidence of downslope creep suggesting deposition on a steep delta front. Sandstone debrites are common, as are tuffaceous beds with evidence of reworking by bottom currents. Sandstone and tuff event beds contain common Skolithos and Arenicolites with less frequent Planolites. Interbedded mudstones contain infrequent Teichichnus.The Caerfai Bay Shales coarsen and thicken-upwards into the Caerbwdy Sandstone. Debrite sandstone beds dominate, and bed thickening upsection indicates progradation of the delta system. The dominant trace fossil within the unit is Teichichnus which is constrained to the top few centimetres of individual sandstone beds. Less common are Planolites and Palaeophycus.The size and abundance of Teichichnus in the Caerfai Group are similar to, or exceed those in contemporaneous sections in Avalonia and Baltica. Local environmental factors were probably the primary control on trace fossil distribution. Substrate and salinity are proposed as the main influences on ichnodiversity and size in the Caerfai Group. Teichichnus is most abundant in the St. Non's Sandstone, where the proximal reaches of the delta experienced reduced salinity and possibly higher oxygen levels. Burrow diameter progressively decreases upsection, and bioturbation becomes less pervasive in subsequent units of the Caerfai Group. This indicates that by the Cambrian Stage 2, significant advances into reduced salinity environments had occurred, along with the dispersal of the Teichichnus producer throughout the Avalonia-Baltica seaway. Correlation within Avalonia indicates that a transgression in the Cambrian Stage 2 initiated the deposition of the Caerfai Group marking the onset of the Welsh Basin.     

Feeding strategy and locomotion of Cambrian hyolithides

The Chengjiang (Cambrian Stage 3) and Balang (Cambrian Stage 4) Konservat-Lagerstätten of South China have produced abundant hyolithide hyoliths; however, little attention has been paid to their feeding strategy and the role it played in the ecosystem. Hyolithides preserved in coprolites from the Chengjiang Biota and associated with a Tuzoia carcass from the Balang Fauna reveal the fluid feces consuming and scavenging strategies of this group. Size distribution of hyolithides demonstrates that their dietary habit is ontogenetically dependent, with juveniles having ingested organic-rich material whereas adult food consumption was more likely by a variety of species-dependent methods The first discovery of hyolithides in association with locomotion traces and burrows indicates they were not only epibenthic vagrants, but also shallow horizontal burrowers. The new discoveries reported herein enhance our understanding of the feeding strategy and other behaviours of Cambrian hyolithides.     

Enigmatic polychaete jaws from the Silurian of Gotland, Sweden

The jaws of the new polychaetaspid polychaete, Oenonites? honki, from the Silurian of Gotland, Sweden, differ from most Palaeozoic polychaete jaws. They exhibit enigmatic microstructural features in that they appear rough and give a corroded, or weathered impression. The altered microstructure of the jaws suggests a jaw chemistry and/or composition differing in some way from that of the co‐occurring polychaete taxa. The jaws appear to have limited preservational potential and/or were particularly susceptible to secondary processes, resulting in microstructural alteration. Commonly, a row of distinct pits occurs on the outer face, especially of the first right maxillae (MIr). Because these pits are interpreted as associated with the dentary, the term ‘denticle marks’ is suggested. The pits may be the result of primary or secondary physical wear, or, more probably, secondary chemical alteration of localized mineral deposits. The primary function of such mineral deposits was to harden those parts of the surface that were exposed to great stress. The restricted occurrence of O.? honki, coupled with occasional increases in abundance (especially in the Halla Formation, unit b), indicates a preference for shallow marine, high‐energy environments, particularly in reefal pockets with calcilutitic sediments. Highest frequency coincides with faunas characteristically containing a few labidognath species also displaying high frequencies.     

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.     

贵州早-中寒武世凯里组类似苔藓植物化石

贵州台江早-中寒武世凯里组保存大量的生物化石,它构成了寒武纪重要的生物群之一--凯里生物群.在凯里生物群中保存有丰富的碳质薄膜藻类化石,新发现类似苔藓植物化石(Parafunaria sinensis Yang(gen.et sp.nov)),它具有藓类植物所拥有的典型的叶状体轮生现象、孢朔、孢朔柄和复杂的根系特征.推测它是藓类植物祖先,它为进一步研究藓类植物和陆生高等植物的起源和演化提供了新的线索.     

贵州早-中寒武世凯里组类似苔藓植物化石(英文)

贵州台江早-中寒武世凯里组保存大量的生物化石,它构成了寒武纪重要的生物群之一——凯里生物群。在凯里生物群中保存有丰富的碳质薄膜藻类化石,新发现类似苔藓植物化石(Parafunaria sinensis Yang(gen.et sp.nov)),它具有藓类植物所拥有的典型的叶状体轮生现象、孢朔、孢朔柄和复杂的根系特征。推测它是藓类植物祖先,它为进一步研究藓类植物和陆生高等植物的起源和演化提供了新的线索。     

Reduction in the number of Early Cambrian helicoplacoid species

Helicoplacoid echinoderms occur in Lower Cambrian shales and mudstones along the northern coast of Laurentia, now western North America. The group originally contained four genera and nine species, diagnosed by differing ambulacra, plate length-to-width ratios, and plate ornament. Ambulacra are identical in all specimens except for the single example of Waucobella nelsoni, and the remaining morphologic differences are caused by ontogeny. The characters used to diagnose seven of the original nine species are therefore invalid, reducing the group to three genera and three species. These are: Helicoplacus gilberti Durham and Caster, 1963, Waucobella nelsoni Durham, 1967, and Polyplacus kilmeri Durham, 1967, which has a test composed of ambulacra, and likely represents a developmental oddity.     

Spirodesmos milanai n. isp.: A Shallow-Water Spiral Trace Fossil from the Cambrian of the Eastern Cordillera,Northwest Argentina

Abstract

A new ichnofossiliferous locality in Salta Province (northwest Argentina) contains an association with numerous irregular spiral traces assigned to Spirodesmos milanai n. isp., in mature sandstones and quartzites with rippled bedding surfaces, rare wavy lamination and cross-bedded stratification. This record of early spiral behavior is interpreted as a primitive grazing method formed on muddy laminae above sand layers, and is related to a feeding strategy of an annelid-type of organism. Associated traces are Cruziana cf. semiplicata, Diplocraterion isp., Monocraterion isp., Rusophycus isp., Skolithos linearis Haldeman and Skolithos magnus Howell. The ichnoassemblage is similar to a shallow-water ichnoassociation from the Permian Ecca Group of South Africa (Mason et al., 1983 Mason, T. R., Stanistreet, I. G. and Tavener-Smith, R. 1983. Spiral trace fossils from the Permian Ecca Group of Zululand. Lethaia, 16: 241–247. [Crossref], [Web of Science ®] , [Google Scholar]).     

A new trace fossil from the Middle Cambrian of the Grand Canyon, Arizona, USA

An unusual trace fossil from the Middle Cambrian Bright Angel Shale Formation of the Grand Canyon, Arizona, consisting of two rows of biserially opposed circular or oval depressions, with alternating symmetry, is described as Bicavichnites martini ichnogen. et ichnosp. nov. While specimens of this trace may represent the surface expression of a complex treptichnid-like burrow system, it is more likely that they are the walking traces of a bilaterally symmetrical benthic animal. The large variation in external width within the trackways is inconsistent with an arthropod producer. The trackway morphology and presence of limb spur marks suggest that the trace maker may have been a lobopodian, a Cambrian relative of modern onychophorans.     

Early Cambrian Salterella and Volborthella (Phylum Agmata) re-evaluated

The genus Volborthella is removed from synonymy under Salterella and placed in the extinct phylum Agmata. Volborthella is more restricted geographically and occurs in older Lower Cambrian strata than Salterella, but distinguishing younger specimens of that genus from poorly preserved Salterella is difficult. Vologinella is not related to Volborthella or other Agmata. Other genera assigned to the phylum by various workers are discussed, but their assignment to this phylum is either enigmatic or rejected. The published concept that specimens of Volborthella and Salterella are agglutinated sclerites or spines on a larger soft-bodied organism is discussed. The material on which this notion is based is re-illustrated and the notion rejected in favor of the view that these conical forms are the hard parts of discrete organisms.     

A spinose stem group brachiopod with pedicle from the Middle Cambrian Burgess Shale

A new genus and species of a Middle Cambrian stem group brachiopod, Acanthotretella spinosa n. gen. and n. sp., is described from the Burgess Shale Formation. Most of the 42 specimens studied came from the Greater Phyllopod bed (Walcott Quarry) and were collected from five bed assemblages, each representing a single obrution event. Specimens are probably preserved within their original habitat. In contrast to all brachiopods known from the Burgess Shale, the shells of the new stem group brachiopod are often deformed and do not show signs of brittle breakage, which suggests that the valves were originally either entirely organic in composition or, more likely, had just a minor mineral component. Acanthotretella spinosa differs from all the other described Cambrian brachiopods in that it is covered by long, slender and possibly partly mineralized spines that are posteriorly inclined at an oblique angle away from the anterior margin. The spines penetrate the shell and are mainly comparable with the thorn‐like organic objects that have been inferred from early siphonotretoid brachiopods. The pedicle was slender and was composed of a central coelomic region and emerged from an apical foramen at the end of an internal pedicle tube. The finding of a pedicle attached to the macrobenthic algae Dictyophycus and other epibenthos implies that A. spinosa did not have an infaunal mode of life. The visceral region and interior characters are poorly preserved.     

Early Cambrian biogeography in the North Atlantic region

A model of Early Cambrian trilobite biogeography in the North Atlantic region (Theokritoff 1979. Lethaia 12 , 281–295) is reviewed and modified in the light of new data. The presence of Wanneria in conglomerates bearing the Elliptocephala asaphoides fauna strengthens correlation of the E. asaphoides fauna as well as, by implication, the upper part of the Calluvia Zone with the middle part of the Bonnia-Olenellus Zone. The faunas of the Bastion Formation of East Greenland and of the Devil's Cove Limestone of northern Newfoundland are reinterpreted as shelf faunas. Additional occurrences of Archaeocyatha are noted in Spain and northwest Scotland but reports of Archaeocyatha in northern Norway and East Germany are not authenticated. Neither Salterella nor 'Volborthella' are now recognized as characteristic of any trilobite province. Although Ordovician acritarch provinces show congruence with trilobite provinces, available data on Early Cambrian plankton are not sufficient to support algal provincialism. Interpretation of the geological record suggests that the Early Cambrian lapetus Ocean was stratified with a well-mixed oxygenated zone overlying a phosphate-rich oxygen minimum zone and that the Elliptocephala asaphoides fauna should be reinterpreted as a warm-water fauna.