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.     

Cambrian stalked echinoderms show unexpected plasticity of arm construction

Feeding arms carrying coelomic extensions of the theca are thought to be unique to crinoids among stemmed echinoderms. However, a new two-armed echinoderm from the earliest Middle Cambrian of Spain displays a highly unexpected morphology. X-ray microtomographic analysis of its arms shows they are polyplated in their proximal part with a dorsal series of uniserial elements enclosing a large coelomic lumen. Distally, the arm transforms into the more standard biserial structure of a blastozoan brachiole. Phylogenetic analysis demonstrates that this taxon lies basal to rhombiferans as sister-group to pleurocystitid and glyptocystitid blastozoans, drawing those clades deep into the Cambrian. We demonstrate that Cambrian echinoderms show surprising variability in the way their appendages are constructed, and that the appendages of at least some blastozoans arose as direct outgrowths of the body in much the same way as the arms of crinoids.     

Deciphering the early evolution of echinoderms with Cambrian fossils

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

Reconsideration of the supposed naraoiid larva from the Early Cambrian Chengjiang Lagerstätte, South China

The Chengjiang Lagerstätte in the Lower Cambrian of South China yields a small, larva‐like arthropod, which was considered to be a protaspis of naraoiids by many authors. The discovery of a large number of well‐preserved specimens from many new localities has allowed the original study to be revised. The relatively large size, stable morphology and unusual structure of the appendages indicate that these specimens represent adults of a new arthropod, Primicaris larvaformis. The larva‐like outline is considered to have arisen by the heterochronic process of progenesis. In addition, this animal displays primitive aspects of bodyplan and limb morphology that suggest a basal position within arachnomorphs, or perhaps even arthropods, and the similarities to the Vendian arthropod‐like animal Parvancorina probably provide an evolutionary link between Vendian forms and Cambrian arthropods.     

Axial patterning of the pentaradial adult echinoderm body plan

Adult echinoderms possess a highly diverged, pentaradial body plan. Developmental mechanisms underlying this body plan are completely unknown, but are critical in understanding how echinoderm pentamery evolved from bilateral ancestors. These mechanisms are difficult to study in indirect-developing species; in this study, we use the direct-developing sea urchin Heliocidaris erythrogramma, whose accelerated adult development can be perturbed by NiCl₂. We introduce a new nomenclature for the adult echinoderm axes to facilitate discussion of the radially symmetric body plan and the events required to pattern it. In sea urchins, the adult oral–aboral axis is often conflated with the long axes of the five rays; we identify these as distinct body axes, the proximodistal (PD). In addition, we define a circular axis, the circumoral (CO), along which the division into five sectors occurs. In NiCl₂-treated larvae, aspects of normal PD pattern were retained, but CO pattern was abolished. Milder treatments resulted in relatively normal juveniles ranging from biradial to decaradial. NiCl₂ treatment had no effect either on mesodermal morphology or on the ectodermal gene expression response to an inductive mesodermal signal. This suggests that the mesoderm does not mediate the disruption of CO patterning by NiCl₂. In contrast, mesodermal signaling may explain the presence of PD pattern in treated larvae. However, variations in appendage pattern suggest that ectodermal signals are also required. We conclude that CO patterning in both germ layers is dependent on ectodermal events and PD patterning is controlled by mutual ectoderm–mesoderm signaling.     

Origins of the other metazoan body plans: the evolution of larval forms

Bilaterian animal body plan origins are not solely about adult forms. Most animals have larvae with body plans, ontogenies and ecologies distinct from adults. There are two primary hypotheses for larval origins. The first hypothesis suggests that the first animals were small pelagic forms similar to modern larvae, with adult bilaterian body plans evolved subsequently. The second hypothesis suggests that adult bilaterian body plans evolved first and that larval body plans arose by interpolation of features into direct-developing ontogenies. The two hypotheses have different consequences for understanding parsimony in evolution of larvae and of developmental genetic mechanisms. If primitive metazoans were like modern larvae and distinct adult forms evolved independently, there should be little commonality of patterning genes among adult body plans. However, sharing of patterning genes is observed. If larvae arose by co-option of adult bilaterian-expressed genes into independently evolved larval forms, larvae may show morphological convergence, but with distinct patterning genes, and this is observed. Thus, comparative studies of gene expression support independent origins of larval features. Precambrian and Cambrian embryonic fossils are also consistent with direct development of the adult as being primitive, with planktonic larvae arising during the Cambrian. Larvae have continued to co-opt genes and evolve new features, allowing study of developmental evolution.     

A new eocrinoid from the Guanshan Biota (Cambrian Series 2, Stage 4), with implication of the development of different attachment modes in early Cambrian

Eocrinoids are scarce in the Guanshan Biota (Cambrian Series 2, Stage 4), Yunnan Province, southwest China. Here, we introduce a new gogiid: Kunmingeocrinus cupuliformis n. gen. n. sp. which is characterized by a short stalk and a well-developed attachment disc. Preservation may indicate a weakly biomineralized body for the new taxon. Morphological similarities between the new taxon and other eocrinoids with attachment discs from Cambrian Lagerstätten of Guizhou Province (Series 2, Stage 4 and Miaolingian, Wuliuan) might suggest a similar mode of life. The eocrinoids from the Guanshan Biota possibly utilized different attachment modes.     

The Cambrian explosion triggered by critical turning point in genome size evolution

The Cambrian explosion is a grand challenge to science today and involves multidisciplinary study. This event is generally believed as a result of genetic innovations, environmental factors and ecological interactions, even though there are many conflicts on nature and timing of metazoan origins. The crux of the matter is that an entire roadmap of the evolution is missing to discern the biological complexity transition and to evaluate the critical role of the Cambrian explosion in the overall evolutionary context. Here, we calculate the time of the Cambrian explosion by a “C-value clock”; our result quite fits the fossil records. We clarify that the intrinsic reason of genome evolution determined the Cambrian explosion. A general formula for evaluating genome size of different species has been found, by which the genome size evolution can be illustrated. The Cambrian explosion, as a major transition of biological complexity, essentially corresponds to a critical turning point in genome size evolution.     

Arthropod body-plan evolution in the Cambrian with an example from anomalocaridid muscle

The ancestor of the arthropods is widely thought to have possessed a hydrostatic skeleton surrounded by peripheral longitudinal and circular musculature, as exhibited by the extant onychophorans. However, the transition to a lever-style musculature system with an articulating exoskeleton poses a difficult problem in functional evolution: did the musculature or the exoskeleton evolve first, and how? Here, by reference to the musculature of the Lower Cambrian stem-group arthropod Pambdelurion, the problem is resolved in terms of preadaptation and functional degeneracy without recourse to saltational notions. Cambrian taxa lying in the stem-groups of the modern phyla may thus be shown to provide unique evidence for the functional progression involved in the assembly of the extant body plans and obviate the need for exotic genetic or developmental mechanisms to explain the evolution of integrated and complex body plans. The notion of the phylum representing a particularly significant level of organization is thereby brought into question.     

【特约论文】寒武纪大爆发的过去、现在与未来

人类探索认识寒武纪大爆发的过程先后经历了神创论、渐变论和爆发式演化思想的影响, 形成了越来越接近真理的重要科学认识。寒武纪大爆发本质上是动物门类的大爆发, 同时伴随着属种多样性的增长、体型增大、生物矿化以及海洋生态系统的重大变革。寒武纪大爆发是多因素制衡的自然历史过程, 任何单因素的内、外因驱动假说都不足以解释寒武纪大爆发的复杂过程。分子遗传基础和宜居环境分别是寒武纪大爆发的内在和外在必要条件, 但不是必要充分条件。因此, 寒武纪大爆发, 与其他所有生命演化事件一样, 必须通过生态作用实现。寒武纪大爆发是在氧气和其他资源不太受限、环境多变、生态荒芜的外在条件下, 基因发育调控系统预先高配的内在条件下, 发生的动物门类大爆发。在埃迪卡拉纪晚期, 多细胞的文德生物与动物采取了不同的体型发育策略和演化途径, 共享海底两千万年后, 文德生物灭绝。此后, 动物在寒武纪最初的两千万年间快速分异演化, 产生了大量演化结局各不相同的动物门类, 生态适应策略同步分化, 形成了以动物为主导的海洋生态系统, 踏上显生宙的演化征程。目前对寒武纪大爆发的探索和研究存在着偏向性或局限性, 主要关注动物门类演化和生存环境氧化还原条件, 没有将海洋生态系统作为统一整体开展全面系统的研究工作。未来探索有必要在全球范围内开展广泛全面的研究工作, 揭示寒武纪大爆发时期海洋生态系统的时空演变。     

Ontogeny,morphology and taxonomy of the soft‐bodied Cambrian ‘mollusc’ Wiwaxia

The soft‐bodied Cambrian organism Wiwaxia poses a taxonomic conundrum. Its imbricated dorsal scleritome suggests a relationship with the polychaete annelid worms, whereas its mouthparts and naked ventral surface invite comparison with the molluscan radula and foot. 476 new and existing specimens from the 505‐Myr‐old Burgess Shale cast fresh light on Wiwaxia's sclerites and scleritome. My observations illuminate the diversity within the genus and demonstrate that Wiwaxia did not undergo discrete moult stages; rather, its scleritome developed gradually, with piecewise addition and replacement of individually secreted sclerites. I recognize a digestive tract and creeping foot in Wiwaxia, solidifying its relationship with the contemporary Odontogriphus. Similarities between the scleritomes of Wiwaxia, halkieriids, Polyplacophora and Aplacophora hint that the taxa are related. A molluscan affinity is robustly established, and Wiwaxia provides a good fossil proxy for the ancestral aculiferan – and perhaps molluscan – body plan.     

Morphology and phylogenetic interpretation of a new Cambrian edrioasteroid (Echinodermata) from Spain

Abstract: The edrioasteroid, Aragocystites belli gen. nov. sp. nov. from the middle Cambrian Murero Formation of Spain, is described based on a small number of very well‐preserved specimens. Important anatomical characteristics include star‐shaped and pseudoclavate theca, rare or absent epispires, well‐developed interradially positioned oral plates and several unorganized cover plates associated with each widely exposed flooring plate. A phylogenetic analysis including several Cambro–Ordovician species shows it is more derived than Stromatocystites and Totiglobus but is a sister group to a clade comprising Cambraster and Edriodiscus. Ontogenetic observations based on juveniles of 5 mm in diameter suggest that this species changed thecal shape markedly during growth. A. belli gen. nov. sp. nov. probably lived in quiet environments where it attached directly to the sea floor on stabilized substrates.     

Carpozoan echinoderms from the Middle Cambrian (Mayaktakh Formation) of Siberia (lower reaches of the Lena River)

Two new monotypic genera and a new monotypic family Rozanovicystidae of the class Cincta are described from the Middle Cambrian (Mayaktakh Formation) of the northeastern part of the Siberian Platform. Based on data obtained by morphofunctional analysis, there is a possibility that these animals could move by jet propulsion by ejecting water from the opercular aperture. Probable homologies of the main skeletal parts are discussed.     

The brachiopod fold: a neglected body plan hypothesis

Attention is drawn to Nielsen's radical body plan concept, here named the 'brachiopod fold hypothesis', under which brachiopods and phoronids are recognized to be transversely folded across the ontogenetic anterior–posterior axis so that, to make useful comparisons with other phyla, these organisms must be conceptually unfolded. Under the hypothesis brachiopod brachial and pedicle shell valves are respectively 'anterior' and 'posterior' rather than 'dorsal' and 'ventral' as traditionally described. The hypothesis makes sense of the symmetry axes of the brachiopod shell, is consistent with various indications from fossil and Recent brachiopods, and gives rise to predicted patterns of axis–determining gene expression that differ from those obtaining under the traditional view of the body plan, whilst the variety of folding movements in different lineages implies that superficially dissimilar morphogenetic folds may be fundamentally homologous. Convergent folding patterns are noted in some other organisms. A previous conjecture that inarticulate linguloid brachiopods were derived from halkieriid–like ancestors is elaborated with proposals that recognize possible functional continuities of coelomic and marginal sclerite functions, and it is noted that an ancestrally facultative fold could have become incorporated by genetic assimilation into the brachiopod developmental program. An experimental approach is outlined to test the possibility that some members of the 'small shelly fauna' may have been members of the halkieriid–like brachiopod stem lineage and it is also suggested that buoyancy modification may have been an important function of mineralization amongst Lower Cambrian floaters and swimmers, since negative buoyancy would facilitate access to the benthic niche.     

Cambrian cinctan echinoderms shed light on feeding in the ancestral deuterostome

Reconstructing the feeding mode of the latest common ancestor of deuterostomes is key to elucidating the early evolution of feeding in chordates and allied phyla; however, it is debated whether the ancestral deuterostome was a tentaculate feeder or a pharyngeal filter feeder. To address this, we evaluated the hydrodynamics of feeding in a group of fossil stem-group echinoderms (cinctans) using computational fluid dynamics. We simulated water flow past three-dimensional digital models of a Cambrian fossil cinctan in a range of possible life positions, adopting both passive tentacular feeding and active pharyngeal filter feeding. The results demonstrate that an orientation with the mouth facing downstream of the current was optimal for drag and lift reduction. Moreover, they show that there was almost no flow to the mouth and associated marginal groove under simulations of passive feeding, whereas considerable flow towards the animal was observed for active feeding, which would have enhanced the transport of suspended particles to the mouth. This strongly suggests that cinctans were active pharyngeal filter feeders, like modern enteropneust hemichordates and urochordates, indicating that the ancestral deuterostome employed a similar feeding strategy.     

The Cambrian evolutionary 'explosion': decoupling cladogenesis from morphological disparity

The origin and differentiation of major clades is often assumed to have occurred in tandem with the 'explosion' of fossil evidence of diverse morphologies ('disparity') at the base of the Cambrian. Evidence is presented that this was not the case. Biogeographical and morphological differentiation among the earliest trilobites reveals incompleteness in the known early Cambrian record; similar evidence can be accrued for other major groups. Phylogenetic analysis proves the likelihood of 'ghost' lineages extending into the Precambrian. The important events in the generation of clades were earlier than the Cambrian 'explosion', at which time the groups become manifest in the fossil record. It is likely that the important phylogenetic changes happened in animals of small size; sister taxa of major groups are shown to be small animals. Decoupling cladogenesis from the Cambrian 'explosion' removes the necessity of invoking unknown evolutionary mechanisms at the base of the Phanerozoic. Genes controlling development may have played a role in generating new morphologies, through heterochrony for example, in the early differentiation of metazoan body plans.