Diverse radial symmetry among the Cambrian Fortunian fossil embryos from northern Sichuan and southern Shaanxi provinces,South China

The Cambrian Fortunian fossil embryos exhibit embryonic development of ancient animals and hence have important bearings on evolutionary developmental biology. They have radial symmetry, and may be early representatives of cnidarians. Here we report new material of three-dimensionally phosphatized fossil embryos from the Fortunian Kuanchuanpu Formation and coeval strata in northern Sichuan and southern Shaanxi provinces, South China. The new material includes previously reported fossil embryos assignable to Pseudooides prima with biradial symmetry or pseudo-hexaradial symmetry, Quadrapyrgites quadratacris with tetraradial symmetry, and Olivooides multisulcatus with pentaradial symmetry. Additionally, we recovered two new types of fossil embryos, i.e., Embryo I with hexaradial symmetry and Embryo II with octaradial symmetry, and they are tentatively suggested to represent new cnidarians. In contrast to the diverse radial symmetry of the Fortunian cnidarians, modern cnidarians exhibit stable tetraradial symmetry in medusozoans, biradial symmetry in anthozoans, and bilateral symmetry in siphonophores (Hydrozoa). The current study supports the view that the tetraradial symmetry of modern medusozoans is a surviving remnant of their Fortunian relatives.     

An intermediate type of medusa from the early Cambrian Kuanchuanpu Formation,South China

The tetraradial or pentaradial fossil embryos and related hatched individuals from the early Cambrian Kuanchuanpu Formation are of great interest for understanding the early evolution of medusozoans. The phylogenetic and evolutionary significance of their external and internal characters (e.g. manubrium, tentacles, septa and claustra) is still controversial. Here we describe a new pentamerous medusozoan, Hanagyroia orientalis gen. et sp. nov., characterized by five well-developed perradial oral lips around a remarkably large manubrium, a conspicuous equatorial groove, and five short interradial pairs of extensile tentacles at the bell margin. Internally, five broad and stout interradial septa join horizontally to form the claustra. Hanagyroia orientalis lacks the frenula, apertural lappet and velarium seen in coeval microfossils and extant cubozoans. Although H. orientalis resembles extant coronate scyphozoans in its round medusa-like bell margin and equatorial groove, cladistic analysis suggests close affinity with cubozoans. Hanagyroia may represent an intermediate morphological type between scyphozoans and cubozoans. The well-developed oral lips and paired short strong tentacles of Hanagyroia suggest direct development.     

Origin and early diversification of the phylum Cnidaria Verrill: major developments in the analysis of the taxon's Proterozoic–Cambrian history

Diploblastic eumetazoans of the phylum Cnidaria originated during the Neoproterozoic Era, possibly during the Cryogenian Period. The oldest known fossil cnidarians occur in strata of Ediacaran age and consist of polypoid forms that were either nonbiomineralizing or weakly so. The oldest possible anthozoans, including the genus Ramitubus, may be related to tabulate corals and occur in the Doushantuo Lagerstätte (upper Doushantuo Formation, South China), the age of which is poorly constrained (approximately 585 Ma?). Conulariid scyphozoans may first appear as early as 635–577 Ma (Lantian Formation, South China). A definite conulariid, most similar to Palaeozoic species assigned to the genus Paraconularia, occurs in association with the possible scyphozoan, Corumbella werneri, in the latest Ediacaran (c. 543 Ma) Tamengo Formation of Brazil. Basal Cambrian (c. 540 Ma) phosphorites in the upper Kuanchuanpu Formation (South China) yield solitary polyps of the oldest probable anthozoan (Eolympia pediculata), which appears to have been a stem hexacorallian. This same formation contains fossils interpreted by some authors as pentaradial cubozoan polyps; however, both the oldest known cubozoans and the oldest hydrozoans, all medusae, may actually occur in the Cambrian (Series 3, c. 505 Ma) Marjum Formation (Utah, USA). Although these recently published palaeontological data tend to corroborate the hypothesis that Cnidaria has a relatively deep Neoproterozoic history, the timing of major internal branching events remains poorly constrained, with, for example, the results of some molecular clock analyses indicating that the two cnidarian subphyla (Anthozoaria and Medusozoa) may have originated as many as one billion years ago. Further progress towards elucidating the evolution and early fossil record of cnidarians may accrue from: (1) an intensive search for phosphatized soft parts in possible anthozoans from the Ediacaran Doushantuo Formation; (2) an expanded search for Ediacaran conulariids; and (3) additional detailed analyses of the taphonomy and preservation of Ediacaran and Cambrian cnidarians, including possible pentaradial cubozoan polyps from the Fortunian upper Kuanchuanpu Formation.     

A first report of Sphenothallus Hall, 1847 in the Cambrian of Variscan Europe

Ten specimens of two phosphatic fossils have been recently discovered in lower and middle portions of Middle Cambrian Jince Formation in the Czech Republic. They are attributed to the genus Sphenothallus Hall, 1847 and described as two separate species; comparatively small conchs are described as S. kozaki sp. nov., the much larger specimens characterized by its smooth and partly flexible organo-phosphatic walls of shell are determined as ?S. kordulei sp. nov. Sphenothallus is known to range from Cambrian to Permian and accommodates numerous species. However, its Cambrian distribution is considerably restricted. Generally rare specimens have been described from Lower to Middle Cambrian of Laurentia and from the Lower Cambrian of Gondwana and peri-Gondwana. The new record of Sphenothallus from the Jince Biota represents a notable extension of their geographic range.     

Conulariid–like fossil from the Vendian of Russia: a metazoan clade across the Proterozoic/Palaeozoic boundary

A conulariid–like fossil is described for the first time from Upper Vendian (late Neoproterozoic) deposits. Vendoconularia triradiata gen. et sp. nov. is found with other fossils in poorly cemented, siliciclastic deposits of the lower part of the Ust'–Pinega Formation, exposed in the middle reaches of the Onega River, north of the Russian Platform. This fossil assemblage, stratigraphically lowermost in the region, includes Swartpuntia Narbonne, endemic Ventogyrus Ivantsov and Grazhdankin, Calyptrina–like tubular forms, and other new, as yet undescribed, species. Vendoconularia is described here using morphological terms developed for conulariids owing to their striking similarity. However, the six faces of the conical test and three–fold symmetry may reflect some phylogenetic connection with the larger clade of Trilobozoa, a cnidarian class that radiated during the Late Vendian–Early Cambrian and then became extinct.     

Asymmetry in an Ordovician conulariid cnidarian

Sendino, C., Zágor?ek, K. & Taylor, P.D. 2012: Asymmetry in an Ordovician conulariid cnidarian. Lethaia, Vol. 45, pp. 423–431. Conulariids are fossils of the presumed polyps of an extinct scyphozoan cnidarian group. Their cone‐shaped skeletons normally show perfect tetraradial symmetry. However, in the Ordovician species Metaconularia anomala ( Barrande 1867 ) from Drabov (Czech Republic), tetraradial symmetry is compromised in three ways: (1) the skeleton often shows torsion; (2) the four sides may vary in width at the same level within one individual; and (3) one side may be deleted to give a triradial skeleton. Almost 2000 specimens were studied in museum collections. About 56% of analysed specimens are twisted in an anticlockwise direction (sinistral) when viewed from the apex towards the aperture, 28% show no torsion, 1% exhibit clockwise torsion (dextral) and the remaining 15% cannot be classified. Maximum measured torsion rate was 1.5°/mm. A significant negative correlation between torsion rate and length suggests that more highly torted individuals may have survived less well. Almost 5% of individuals show loss of one side for at least part of their lengths. Although many individuals have four sides of equal width, in a significant proportion the sides are of unequal width, up to a maximum/minimum side width ratio of 2 (i.e. widest face twice the width of the narrowest). In the absence of a satisfactory taphonomic model to explain the asymmetries, they are regarded as mirroring asymmetries in the living conulariids, with the strong preference for sinistral torsion interpreted as an example of a fixed asymmetry that was genetically controlled and heritable. It is speculated that the signalling protein Nodal as well as Hox‐like genes were involved in controlling the asymmetries described in M. anomala.     

Tiny sea anemone from the Lower Cambrian of China

Background

Abundant fossils from the Ediacaran and Cambrian showing cnidarian grade grossly suggest that cnidarian diversification occurred earlier than that of other eumetazoans. However, fossils of possible soft-bodied polyps are scanty and modern corals are dated back only to the Middle Triassic, although molecular phylogenetic results support the idea that anthozoans represent the first major branch of the Cnidaria. Because of difficulties in taxonomic assignments owing to imperfect preservation of fossil cnidarian candidates, little is known about forms ancestral to those of living groups.

Methods and Findings

We have analyzed the soft-bodied polypoid microfossils Eolympia pediculata gen. et sp. nov. from the lowest Cambrian Kuanchuanpu Formation in southern China by scanning electron microscopy and computer-aided microtomography after isolating fossils from sedimentary rocks by acetic acid maceration. The fossils, about a half mm in body size, are preserved with 18 mesenteries including directives bilaterally arranged, 18 tentacles and a stalk-like pedicle. The pedicle suggests a sexual life cycle, while asexual reproduction by transverse fission also is inferred by circumferential grooves on the body column.

Conclusions

The features found in the present fossils fall within the morphological spectrum of modern Hexacorallia excluding Ceriantharia, and thus Eolympia pediculata could be a stem member for this group. The fossils also demonstrate that basic features characterizing modern hexacorallians such as bilateral symmetry and the reproductive system have deep roots in the Early Cambrian.     

Occurrence ofByronia Matthew andSphenothallus Hall in the Lower Cambrian of China

Rare phosphatic tubular fossils from the Lower-Middle Cambrian Kaili Formation of Guizhou Province, southern China were originally identified as non-calcified algae or ‘worms’ (ScoleciellusLiu). Re-examination of these fossils indicates that specimens identified as non-calcified algae areSphenothallus taijiangensis n. sp., while specimens identified asScoleciellus belong toByronia natus (Liu).Sphenothallus taijiangensis andByronia natus from Lower Cambrian strata in the Kaili Formation are the oldest known representatives of their genera. In addition,B. natus (Liu) is the only known species ofByronia with the exception ofB. annulata Matthew (Middle Cambrian, British Columbia). CambrovitusMao et al., a tubular fossil from Middle Cambrian strata in the Kaili Formation, originally was classified as a hyolithid. However, the discovery of a nearly complete specimen possessing an apical attachment disk shows thatCambrovitus, likeByronia andSphenothallus, probably was a thecate cnidarian polyp.     

Anatomy and phylogenetic significance of Eoconularia loculata, a conulariid from the Silurian of Gotland

The type material of Eoconularia loculata (Wiman, 1895), a conulariid with high, bifurcated septa, was originally found in an erratic boulder, hence the source-rock is as yet unidentified. Recently, a rich material of the species has been discovered at eleven localities in the Silurian Hemse Beds of Gotland Another two localities in the Slite Beds (Wenlockan, Sheinwoodian) revealed what is assumed to be the ancestor of E loculata. This ancestral form probably constitutes a separate taxon distinguished from E. loculata in having simple, unbranched septa. E. loculata is re-described, and four ontogenetic stages in septal development are recognized. During stage 1, the most juvenile stage, the septa are simple. The septa in stages 3 and 4, the adult stages, are coarse and bifurcated. The affinities of conulariids are discussed, with the conclusion that the group shares a number of similarities with modem scyphomans. The microstructures of the exoskeleton show several similarities with coronatids, and the septa are interpreted to be homologous with the internal structures of stauromedusids. The stratigraphical distribution of all currently known septate conulariids suggests that septa were a primitive morphologic feature ranging from early? Ordovician to late Silurian. The simplest type, however, persisted at least into the early Permian. Five of the eleven described septa types have been found only among the conulariids from Gotland. □Conulariida, EOCONULARIA LOCULATA, bifurcated septa, taxonomy, ecology, morphology, Scyphozoa, Sweden, Gotland, Silurian.     

Early Cambrian Pentamerous Cubozoan Embryos from South China

Background

Extant cubozoans are voracious predators characterized by their square shape, four evenly spaced outstretched tentacles and well-developed eyes. A few cubozoan fossils are known from the Middle Cambrian Marjum Formation of Utah and the well-known Carboniferous Mazon Creek Formation of Illinois. Undisputed cubozoan fossils were previously unknown from the early Cambrian; by that time probably all representatives of the living marine phyla, especially those of basal animals, should have evolved.

Methods

Microscopic fossils were recovered from a phosphatic limestone in the Lower Cambrian Kuanchuanpu Formation of South China using traditional acetic-acid maceration. Seven of the pre-hatched pentamerous cubozoan embryos, each of which bears five pairs of subumbrellar tentacle buds, were analyzed in detail through computed microtomography (Micro-CT) and scanning electron microscopy (SEM) without coating.

Results

The figured microscopic fossils are unequivocal pre-hatching embryos based on their spherical fertilization envelope and the enclosed soft-tissue that has preserved key anatomical features arranged in perfect pentaradial symmetry, allowing detailed comparison with modern cnidarians, especially medusozoans. A combination of features, such as the claustrum, gonad-lamella, suspensorium and velarium suspended by the frenula, occur exclusively in the gastrovascular system of extant cubozoans, indicating a cubozoan affinity for these fossils. Additionally, the interior anatomy of these embryonic cubozoan fossils unprecedentedly exhibits the development of many new septum-derived lamellae and well-partitioned gastric pockets unknown in living cubozoans, implying that ancestral cubozoans had already evolved highly specialized structures displaying unexpected complexity at the dawn of the Cambrian. The well-developed endodermic lamellae and gastric pockets developed in the late embryonic stages of these cubozoan fossils are comparable with extant pelagic juvenile cubomedusae rather than sessile cubopolyps, whcih indicates a direct development in these fossil taxa, lacking characteristic stages of a typical cnidarian metagenesis such as planktonic planula and sessile polyps.     

Eoandromeda and the origin of Ctenophora

The Ediacaran fossil Eoandromeda octobrachiata had a high conical body with eight arms in helicospiral arrangement along the flanks. The arms carried transverse bands proposed to be homologous to ctenophore ctenes (comb plates). Eoandromeda is interpreted as an early stem‐group ctenophore, characterized by the synapomorphies ctenes, comb rows, and octoradial symmetry but lacking crown‐group synapomorphies such as tentacles, statoliths, polar fields, and biradial symmetry. It probably had a pelagic mode of life. The early appearance in the fossil record of octoradial ctenophores is most consistent with the Planulozoa hypothesis (Ctenophora is the sister group of Cnidaria + Bilateria) of metazoan phylogeny.     

华北寒武系第二统辛集组软舌螺动物化石群*

软舌螺动物是华北板块寒武系小壳化石基本组成之一,也是寒武纪生命大爆发在华北地区的重要见证者。本文系统描述了华北板块南缘-西南缘(河南-陕西地区)寒武系第二统辛集组软舌螺化石9属14种及1未定种。包括直管螺类6属9种:Conotheca australiensis、Cupitheca holocyclata、Cupitheca costellata、Cupitheca? decollata、Tegminites hymenodes、Triplicatella disdoma、Triplicatella xinjia、Protomicrocornus triplicensis和Longxiantheca mira;软舌螺类3属5种及1未定种:Microcornus eximius、Microcornus petilus、Parkula cf. esmeraldina、Parkula bounites、Parakorilithes mammillatus和Parakorilithes sp.。基于大量保存的精美聚合标本,本文准确重建了锥壳和口盖在软舌螺不同属种中的配置。此外,新材料显示华北辛...     

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

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

山东潍坊寒武系苗岭统乌溜阶馒头组上部的腕足动物

寒武系腕足动物属种多样性高、个体数量丰富、形态差异明显、地理分布广泛,具有辅助寒武系三叶虫生物地层划分和对比的潜力.华北板块寒武系苗岭统沉积和化石记录发育良好,是中国苗岭统的经典研究区之一.前人己针对华北寒武系苗岭统乌溜阶腕足动物的系统古生物学开展了一系列基础工作,但这些相关研究主要集中于辽宁地区,目前对华北其他地区苗...     

Graptolite (Hemichordata,Pterobranchia) preservation and identification in the Cambrian Series 3

Due to inadequate preservation, pterobranchs are often difficult to identify in the fossil record, and a better understanding of preservational modes and diagenetic and metamorphic effects is needed for their recognition. Pterobranch hemichordates are common in Cambrian Stage 5 and younger sedimentary rocks, but are frequently overlooked. Often, pterobranch hemichordate colonies have been considered to be algal remains or hydroids. Re‐examination of Cambrian Burgess Shale algae reveals that the genera Yuknessia and Dalyia can be recognized as putative early representatives of pterobranch hemichordates. Distinct fusellar construction of the individual zooidal tubes and branching of the creeping proximal part of the colonies are found in the morphologically similar rhabdopleurid pterobranch genus Sphenoecium. The erect tubes of Sphenoecium do not branch and can reach a length of several centimetres. The development of the fusellar construction in this taxon shows a highly irregular development of the suture patterns, but a fairly consistent height of the individual fuselli. The taxon is widely distributed in the Cambrian Series 3, but has regularly been identified as a hydroid or an alga. Sphenoecium wheelerensis from the Cambrian Wheeler Shale of Utah is described as new.     

Giving the early fossil record of sponges a squeeze

Twenty candidate fossils with claim to be the oldest representative of the Phylum Porifera have been re‐analysed. Three criteria are used to assess each candidate: (i) the diagnostic criteria needed to categorize sponges in the fossil record; (ii) the presence, or absence, of such diagnostic features in the putative poriferan fossils; and (iii) the age constraints for the candidate fossils. All three criteria are critical to the correct interpretation of any fossil and its placement within an evolutionary context. Our analysis shows that no Precambrian fossil candidate yet satisfies all three of these criteria to be a reliable sponge fossil. The oldest widely accepted candidate, Mongolian silica hexacts from c. 545 million years ago (Ma), are here shown to be cruciform arsenopyrite crystals. The oldest reliable sponge remains are siliceous spicules from the basal Cambrian (Protohertzina anabarica Zone) Soltanieh Formation, Iran, which are described and analysed here in detail for the first time. Extensive archaeocyathan sponge reefs emerge and radiate as late as the middle of the Fortunian Stage of the Cambrian and demonstrate a gradual assembly of their skeletal structure through this time coincident with the evolution of other metazoan groups. Since the Porifera are basal in the Metazoa, their presence within the late Proterozoic has been widely anticipated. Molecular clock calibration for the earliest Porifera and Metazoa should now be based on the Iranian hexactinellid material dated to c. 535 Ma. The earliest convincing fossil sponge remains appeared at around the time of the Precambrian‐Cambrian boundary, associated with the great radiation events of that interval.     

Occurrence of the earliest known Sphenothallus Hall in the Lower Cambrian of Southern Shaanxi Province, China

Three-dimensionally preserved specimens of an indeterminate species of Sphenothallus Hall occur in limestones in the uppermost Guojiaba and lower Xiannudong formations (Qiongzhusian and early Canglangpuan stages, Lower Cambrian) in southern Shaanxi Province, China. The specimens consist of a small (up to 3 mm long), gradually tapered, subcircular to elliptic (transversely) tube that is broken at both ends, commonly with the fracture edge at the wide (apertural) end of each face forming a V- or U-shaped notch. Some specimens preserve the subconical upper (proximal) part of the holdfast, and all specimens are black. Combined scanning and transmission electron microscopy and energy dispersive elemental analyses show that the tube wall is finely laminated, with individual laminae measuring about 1 μm thick, and consists mainly of calcium phosphate but also contains organic material. Sphenothallus specimens from the Lower Cambrian Guojiaba and Xiannudong formations represent the earliest known record of this genus, and indicate that the origin of this taxon is coeval with and possibly related to the Early Cambrian bioradiation event.     

Exceptionally preserved conulariids and an edrioasteroid from the Hunsrück Slate (Lower Devonian,SW Germany)

Abstract: Nineteen partial specimens of Conularia sp., together with an articulated agelacrinitid edrioasteroid and several discinid brachiopods, occur in close association with a probable biological substrate on a small slab of silty Hunsrück Slate (Lower Devonian, Emsian) from Bundenbach, Germany. Most of the conulariids occur in V‐like pairs or in a single cluster of 12 specimens arranged in a fan‐like radial pattern. Together with the edrioasteroid and (possibly) brachiopods, the conulariids probably were attached to the substrate in life and then were buried and possibly killed by a single influx of silty mud. The apertural end of many of the conulariids is partially covered by inwardly folded short lappets, which may have closed in response to rapid (but gentle) burial. Rock matrix in the apertural region of the peridermal cavity of nearly all of the conulariids exhibits irregular, variably dense concentrations of pyrite. The concentrations occur almost exclusively within the conulariids, where they probably formed as a result of the decay of retracted conulariid soft parts. Although the concentrations lack clearly defined anatomical features that can be unambiguously homologized with particular anatomical structures of any extant taxon, their form and distribution within the conulariids are consistent with the hypothesis that conulariids were polypoid scyphozoans.     

Possible ctenophoran affinities of the Precambrian "sea-pen" Rangea

The Namibian Kuibis Quartzite fossils of Rangea are preserved three-dimensionally owing to incomplete collapse of the soft tissues under the load of instantaneously deposited sand. The process of fossilization did not reproduce the original external morphology of the organism but rather the inner surface of collapsed organs, presumably a system of sacs connected by a medial canal. The body of Rangea had tetraradial symmetry, a body plan shared also by the White Sea Russian fossil Bomakellia and possibly some other Precambrian frond-like fossils. They all had a complex internal anatomy, smooth surface of the body, and radial membranes, making their alleged colonial nature unlikely. Despite a different style of preservation, the Middle Cambrian Burgess Shale frond-like Thaumaptilon shows several anatomical similarities to Rangea. The body plan of the Burgess Shale ctenophore Fasciculus, with its numerous, pinnately arranged comb organs, is in many respects transitional between Thaumaptilon and the Early Cambrian ctenophore Maotianoascus from the Chengjiang fauna of South China. It is proposed that the irregularly distributed dark spots on the fusiform units of the petaloid of Thaumaptilon represent a kind of macrocilia and that the units are homologous with the ctenophoran comb organs. These superficial structures were underlain by the complex serial organs, well represented in the fossils of Rangea. The Precambrian "sea-pens" were thus probably sedentary ancestors of the ctenophores.     

Probable advanced hydroid from the Early Ordovician of China

A fossil of a colonial organism with pyritized soft tissues of elongated fusiform zooids, found in the middle part of the early Floian Fenxiang Formation in Hubei Province of China, probably represents the oldest reliable record of a hydroid cnidarian. The preservation of the fossil is sufficiently different from that of associated carbonized skeletons of graptolites to exclude affinities with this group. The fossil is unlikely to be a bryozoan because of the mode of budding from proximal, not distal, parts of parent zooids, which is typical rather of hydroids. Although no thecae are preserved, the fossil, named Sinobryon elongatum gen. et sp. nov., is suggested to be a thecate hydroid, possibly related to the Haleciidae. The apparent presence of an advanced member of the thecaphoran Macrocolonia clade in strata 470 Ma old means that much of the hydroid (and cnidarian) diversification preceded the Middle Ordovician.