Modelling the Transport of Nutrients in Early Animals

The single-celled ancestors of multi-cellular animals (metazoans) did not need to transport nutrients between cells, but this ability is vital for modern animals. How could intercellular nutrient transport have begun? And how did this influence the early evolution of animals? In this hypothesis, I suggest that nutrients could have passed directly between the cytoplasm of conjoined cells in early compacted cell-balls, along the plane of the closed epithelium. This would have limited early animals to the size and form of modern embryos. The mechanisms that indirectly transport nutrients between discrete cells, via the extracellular fluid within the body-space, are modelled to have evolved sequentially; so comparison of nutrient transport processes could provide evidence of any early divergences of phyla. When the last of the indirect intercellular transport processes for essential nutrients had been developed, the extracellular fluid within the body-space would have contained all necessary nutrients. Then the epithelium could have greatly expanded, and cells lived and divided within the body-space. This development of nutrient transport processes would have enabled animals to greatly increase in size and complexity.     

Trilobite faunal successions across the base of the Furongian Series in the Taebaek Group, Taebaeksan Basin, Korea

The base of the Furongian Series in the Sino-Korean Block has not been clearly defined due to the lack of the index taxon, Glyptagnostus reticulatus. The Sesong Formation of the Taebaek Group, Taebaeksan Basin, Korea, has been known to range from the Guzhangian Stage of the Cambrian Series 3 to the middle Furongian Series, hence embracing the base of the Furongian Series. Silicified polymerid trilobites were recovered from the middle part of the Sesong Formation. Described are a total of 18 polymerid species of 13 genera: Neodrepanura sp. 1, Teinistion sp. 1, Huzhuia sp. 1, Huzhuia sp. 2, Liostracina simesi, Liostracina sp. 1, Parachangshania monkei, Parachangshania rectangularis nov. sp., Placosema bigranulosum, Fenghuangella laevis nov. sp., Baikadamaspis jikdongensis nov. sp., Baikadamaspis sp. 1, Prochuangia mansuyi, Maladioides coreanicus, Alataspis sesongensis nov. gen., nov. sp., Chuangia sp. 1, and ceratopygids genus and species indeterminate 1 and 2. The stratigraphic occurrence of these trilobites provides a basis for recognition of five zones across the base of the Furongian Series (in ascending order): the Neodrepanura, Liostracina simesi, Fenghuangella laevis, Prochuangia mansuyi, and Chuangia zones. The Neodrepanura and Chuangia zones are provisionally adopted from the previous biostratigraphic scheme, while the three other ones are newly proposed. The recommended base of the Furongian Series in the Taebaek Group of Korea coincides with the base of the Fenghuangella laevis Zone, which appears to represent an episode of profound trilobite faunal turnover.     

Proposed GSSP for the base of Cambrian Stage 7, coinciding with the first appearance of Lejopyge laevigata, Hunan, China

The Luoyixi section, exposed in a roadcut along the Youshui River (Fengtan Reservoir), Guzhang County, Hunan Province, China, is proposed as the stratotype for the base of an unnamed stage boundary (base of the Cambrian stage provisionally termed Stage 7). The proposed position of the GSSP is 121.3 m above the base of the Huaqiao Formation, at a horizon coinciding with the first appearance of the cosmopolitan agnostoid trilobite Lejopyge laevigata. The section fulfills all the requirements for a GSSP, and the horizon can be constrained not only with the primary stratigraphic marker (L. laevigata) but also with secondary biostratigraphic, sequence-stratigraphic, and chemostratigraphic correlation tools. The first appearance of L. laevigata is one of the most readily recognizable levels in the Cambrian, and can be correlated with precision to all paleocontinents.     

拟浒苔在澄江生物群中的发现及其生态学意义

报道了早寒武世澄江生物群中的一种新的化石藻类－肠拟浒苔（Enteromophites intestinalis sp.nov）,丰富了对该时期生物多样性的认识,通过现生浒苔属和石拟苔属藻类形态学的研究表明,这两个属在系统演化上可能有密切的亲缘关系,并阐述了澄江生物群可能发生了一个海水环境。     

Evolutionary uniformitarianism

I present a new compilation of the distribution of the temporal distribution of new morphologies of marine invertebrates associated with the Ediacaran–Cambrian (578–510 Ma) diversification of Metazoa. Combining this data with previous work on the hierarchical structure of gene regulatory networks, I argue that the distribution of morphologies may be, in part, a record of the time-asymmetric generation of variation. Evolution has been implicitly viewed as a uniformitarian process where the rates may vary but the underlying processes, including the types of variation, are essentially invariant through time. Recent studies demonstrate that this uniformitarian assumption is false, suggesting that the types of variation may vary through time.     

Ultrastructure of some Cambrian palynomorphs from the Bright Angel Shale, Arizona, USA

Palynomorphs with complex resistant walls have been recovered from several Cambrian deposits in the continental United States. Those recovered from the Bright Angel Shale of Arizona typically preserve both a primary wall, and an outer envelope (synoecosporal wall) that encloses multiple spore-like bodies within. At least three distinct types of primary walls, are recognized with the TEM: 1) a unilaminate wall with a smooth inner surface and a sculptured outer surface, 2) a wall of three unornamented laminae of very uniform thickness, and, 3) a thicker wall with multiple thin, lightly-staining layers embedded in a darker matrix. This third type of primary wall bears a strong resemblance to those of certain Lower Devonian hilate cryptospore monads from the Welsh Borderlands. No extant algae produce spores with walls as thick or as complex, suggesting that these Cambrian palynomorphs were the desiccation-resistant spores of cryptogams belonging to the charophyte–embryophyte lineage. Multilaminate spore walls, which are characteristic of some extant liverworts and Paleozoic cryptospores, may have evolved via the fusion of separate, multiple laminae. This appears to be the primitive plant sporoderm type, but it may have evolved asynchronously with respect to the evolution of the embryophytic development of the sporophyte in land plants.     

A scratch circle origin for the medusoid fossil Kullingia

Kullingia is considered a key taxon in demonstrating the presence of terminal Proterozoic-early Cambrian chondrophorine hydrozoans. However, Kullingia concentrica from the Lower Cambrian of northern Sweden possesses several features that show that it is not a body fossil but that it was formed by current or wave-induced rotation of an anchored tubular organism, possibly a sabelliditid. A scratch circle interpretation applies also to several other reports of Lower Cambrian Kullingia , including Kullingia delicata from the Chapel Island Formation of Newfoundland. Given the considerable number of problematic fossils that have been interpreted as chondrophorines, it is difficult to put an age on the oldest fossil chondrophorines, but it may be as late as the Devonian. Overall, scratch circles appear to be rarely preserved. The occurrence of these scratch circles in Lower Paleozoic storm deposits is probably related to low levels of bioturbation that enhanced both the likelihood of formation and preservation of these structures.     

The soft anatomy of Isoxys minor from the Guanshan fauna,lower Cambrian of Southwest China

Isoxys is a very common Cambrian bivalved arthropod, specimens of which are normally preserved only as valves. The discovery of the soft anatomy of Isoxys may greatly assist understanding affinities and functional morphology. Isoxys minor Luo and Hu in Luo et al., 2008 is the most common representative of all animal species known from the lower Cambrian Guanshan fauna (Cambrian Series 2, Stage 4) at the Shitangshan Section, near Kunming, Yunnan Province, Southwest China. Here we describe and reconstruct the morphology of I. minor on the basis of newly illustrated fossils and a few new specimens that bear soft-parts including new discovery of frontal grasping appendages. Like the soft anatomy of other known Isoxys, it bears a pair of large stalked eyes, a pair of specialized frontal grasping appendages, approximately 12–14 paired biramous limbs, and a helm-like tail exposed outside the valves.     

Revision of Echmatocrinus from the Middle Cambrian Burgess Shale of British Columbia

Echmatocrinus from the Middle Cambrian Burgess Shale of British Columbia was originally described as the earliest crinoid(?) known from the fossil record. Recently, Conway Morris and Ausich & Babcock have questioned whether Echmatocrinus is in fact an echinoderm, comparing it instead to cnidarians with a polyp-like body and pinnate tentacles, and other authors are beginning to use this reinterpretation. We studied the well-preserved holotype of Echmatocrinus brachiatus, two paratypes, and 18 new specimens recovered from different levels in the Burgess Shale sequence at three localities. All are preserved as pyrite films in dark shale with relatively little relief, suggesting a lightly skeletized body. Complete specimens have a long, slightly tapering, large-plated attachment stalk, a conical cup or calyx with numerous small to medium-sized irregular plates, and 7–10 short arms with heavier plating and (in the holotype) soft appendages alternating from opposite sides of several arms. Several morphologic features indicate that Echmatocrinus is an echinoderm and has crinoid affinities: (1) Sutured plates, shown by darker depressed sutures, slightly raised plate centers, and oriented plate ornament, cover all major parts of the body; (2) reticulate surface ornament in the pyrite film on the plates of all specimens matches the ornament in the Burgess Shale edrioasteroid Walcottidiscus, an undoubted echinoderm, but not the pyritized surfaces of other metazoans in the fauna; (3) this distinctive ornament may represent the surface expression of microporous stereom; (4) possible ligament or muscle pads are present between the arm ossicles to fold and unfurl the more heavily plated arms. Within the echinoderms, only crinoids commonly have a calyx attached by a stalk or stem to the substrate and bear erect, moveable, uniserial arms for feeding. Although Echmatocrinus shows some resemblance to octocorals in overall body shape as an attached suspension feeder, almost all the details are different, indicating that Echmatocrinus is most likely unrelated to this group. All complete specimens of Echmatocrinus are attached to hard substrates, either another fossil or skeletal debris. The new specimens indicate that Echmatocrinus was twice as common (about 0.02%) in the Burgess Shale fauna as previously recorded and represents one of the earliest attached, medium-level, skeletized, suspension feeders or microcarnivores in the fossil record.     

湖南花垣中寒武世晚期至晚寒武世早期球接子类

在考察、实测了湘西地区数条剖面后,主要研究了花垣排碧剖面。建立了中寒武统上部至上寒武统下部的球接子类三叶虫带,这在湘西黔东地区尚属首次,为本区与国内、外有关地区的地层对比,提供了直接证据。本文详细讨论了这些对比证据,并系统描述了球接子类三叶虫新种和属于我国首次报道的种。此外,基于微相分析,概述了排碧地区中寒武世晚期至晚寒武世早期的沉积环境。     

Crypto-helical body plan in partially disarticulated gogiids from the Cambrian of South China

All living echinoderms have a pentaradial symmetry that is unique within the Bilateria. However, the Cambrian origin of echinoderm radial/pentaradiate symmetry is a long-standing problem. During the Cambrian (542–488 Ma), gogiids were the most common stalked echinoderm characterized by an “irregularly” plated body. Based on recently discovered material from the Balang Formation (Cambrian Series 2), eastern Guizhou, China, three unusual, partially disarticulated specimens of Guizhoueocrinus have clear evidence for a helical body plan. This helical plating is only evident in partially disarticulated specimens, thus a crypto-helical body construction is present. Crypto-helical construction in a gogiid raises the possibility of a phylogenetic connection among helicoplacoids, gogiids, and Helicocystis. The crypto-helical body construction may be an important evolutionary innovation among pre-radiate echinoderms.     

陕西韩城寒武系张夏组和三山子组三叶虫动物群

陕西韩城寒武系出露良好,化石丰富,从下寒武统上部至奥陶系为连续沉积,底部平行不整合覆盖厚约20m的霍山组石英砂岩之上。本区寒武 系主要以紫色和黄色泥页岩、泥灰岩、灰岩和白云质灰岩为主,张夏组主要以鲕状灰岩为主,夹生物碎屑灰岩,三叶虫主要有Changqingia chalcon,Changqingia luia sp.nov.,Manchuriella macar,Lianglangshania hueir ensis,Crepicephalia convexus,Eilura quadrata,Eilura(?)hanchengensis sp.nov.,Anmocarella chinensis,Dorypyge pergranosa,Dorypyge richthofeni,Liopeishania lubrica,Liopeishania marginata和Damesella paronai等。三山子组以白云质灰岩和灰质白云岩为主,在底部的白云质灰岩中产三叶虫Blackwelderia sp.,Damesops convexus和Cyclolorenzella acalle等,其中、上部未采到化石。     

Benthic primary producers in exceptionally preserved Cambrian biotas of North China

Strata situated within the South China Craton host several key Cambrian macroalgal assemblages, including those of the Chengjiang, Kaili, and Tsinghsutung Lagerstätte. In this report, Cambrian macroalgae are described for the first time from the North China Craton, an area that, during the early Paleozoic, was not contiguous with the South China Craton. The material collected from the Mantou Formation and overlying Zhangxia Formation (Miaolingian Series) in Shandong Province is preserved as carbonaceous compressions and includes abundant simple tubular forms. These fall into two groupings on the basis of size and have characteristics broadly comparable to Fuxianospira gyrata and Sinocylindra yunnanensis, both of which are well-known from the Chengjiang biota and may represent simple siphonous green algae. Additional material includes irregular circular to oval films referable to Morania fragmenta and a tuft-like form superficially similar to Marpolia spissa, taxa with possible cyanophyte affinities described initially on the basis of material from the Burgess Shale but also known from the Cambrian of South China. The material is typical of the “Cambrian Flora” and provides further support for the cosmopolitan distribution of this noncalcified macroalgal flora at low latitude settings during the Cambrian.     

A Cambrian tommotiid preserving soft tissues reveals the metameric ancestry of lophophorates

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Redescription and systematic position of Diandongia pista from the Chengjiang Lagerstätte

Diandongia pista Rong is the thickest-shelled and strongly mineralized of all the brachiopods that have been recorded from the Chengjiang Fauna. It is also one of the most common species in the biota. Previous accounts have concentrated largely on the exceptionally preserved soft-bodied anatomy of this species. Here its detailed internal and external shell morphologies are described for the first time, based on BSEM and SEM observations. Its general shell morphology, pseudointerareas, and pustulose ornamentation are most similar to those of the Botsfordiidae, where many species have a Diandongia-like rhombic pattern of pustules in the juvenile apical region only. On the other hand, other aspects of the muscle pattern and the partly smooth adult ornamentation of D. pista is more suggestive of species within the Neobolidae, like Edreja. The mantle canal system in sub-adults of D. pista is comparable to that of the Eoobolidae and Obolidae, whereas the adult vascular system branches peripherally in dichotomy. D. pista may be the earliest representatives possessing some plesiomorphic characters of the Botsfordiidae.     

贵州寒武纪海绵动物化石组合特征

贵州寒武纪海绵化石丰富。分布广泛,主要产于早寒武世梅树村期的戈仲伍组或牛蹄塘组底部、筇竹寺期的牛蹄塘组和早、中寒武世凯里组。梅树村期的海绵属种单调,主要为六射海绵类麦粒海绵（Triticispongia sp．）及一些海绵骨针,代表寒武纪海绵首次爆发或辐射的前奏和先驱。牛蹄塘组海绵多达13个属,是寒武纪海绵辐射的重要窗口,以普通海绵略占优势,Leptomitus为优势属。凯里组的海绵共计7属,以普通海绵为主。贵州寒武纪海绵动物群的发展具有明显的阶段性和组成的差异性,其演变过程受到环境的制约。贵州早期海绵动物的发现和研究不仅改变了寒武纪生物多样性的认识,而且还为探讨海绵动物本身的起源、辐射提供了重要材料。     

Palaeoecology and evolutionary implications of enrolled trilobites from the Kushan Formation,Guzhangian of North China

Damesella paronai is the earliest enrolled odontopleurid trilobite to be found in the fossil record. Whereas its interlocking devices were very poorly developed, its trunk shows advanced articulations, which only lack an articulating facet, though an anterior notch is present in the outer part of each pleura. Its body pattern and structures did not allow it an encapsulated enrolment style. On the other hand, it is coeval with the early asaphid Monkaspis daulis, found in the same beds and at the same level. This co-inhabiting trilobite had a more advanced structure that enabled it to achieve a fully enrolled, encapsulated style. These superior structures enhance the preservation of enrolled specimens of M. daulis, and it seems to be generally the case that more elegant structures actually improve preservation. The evolutionary trends of the lineages of these two trilobites show that while odontopleurids were very conservative, using the same enrolment style throughout their history, the asaphids developed different enrolment styles, which was one factor in giving them a capacity to occupy different ecological niches and a greater range of environments.     

Discovery of Enteromophites in the Chengjiang Biota and Its Ecological Significance (in English)

The paper deals with a new species of megaalgal fossil— Enteromophites intestinalis sp. nov. found in Chengjiang Biota in the early Cambrian. A morphological comparison of the living Enteromorpha and fossil Enteromophites indicates that there probably is a close relationship between these two genera. Based on a study on the surroundings of living Enteromorpha, Enteromophites intestinalis sp. nov. might have lived in the environment of sea or salt water.     

Unlocking the early fossil record of the arthropod central nervous system

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